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Flowering Plants 

Vol. 10, part 4 















Aclinidiaceac s.s. 









7: ] 












Alismaiaccae 5: 























Pentaphragm atace ac 



4: 69, 593; 6: 




4: 377; 


























Aponogctonaceae 4 

I: 11; 











: 1, 



6: 469, 










8: 549; 












































6: 943; 



















Betulaccae 5: 












Bignoniaceae 8: 












Bixaceae s.s. 











5: : 






4: 13 


; 9: 


6: 949; 









5: 209, 








6:917; 7: 








5: 429; 






5: 1 




























6: 952; 7: 828; 








6: 107, 














8: 1; 



Scyphosiegiaceac 5 


'; 6: 


Cappar(id)accae 6:61; 




7: 1; 














4: 175; 6 


; 9: 





4: 280, 















Cclastraceae 6: 227, : 






































4:99,594; 6: 




































8: 205; 













6: 293, 







4:533; 5: 








Conifc rales 






4:349; 6: 

: 1, 











5:495; 6: 











4: : 









5: 558; 6: 936; 7 


; 9: 

















Corynocarpaceae 4 


; 5: 










Crassulaceae 4 

: 197 

; 9: 
























5:555; 6:983; 



; 9: 

















Violaceae 7: 179, 




Cypcraccae 7: 








Xyridaccae 4: 366, 







































SERIES I ..^..^^..^ . VOLUME 10 



Dr. C. G. G. J. VAN STEENIS (f 1986) 
Dr. W. J. J. O. DE WILDE 


1 9 K 4 I y « y 

Library in Congress Catalog Card Number 72-175112 
ISBN 0-7923-0421-7 

Published by Kluwer Academic Publishers, 

P.O. Box 17, 3300 AA Dordrecht, The Netherlands. 

Kluwer Academic Publishers incorporates 

the publishing programmes of 

D. Reidel, Martinus Nijhoff, Dr W. Junk en MTP Press. 

Sold and distributed in the U.S.A. and Canada 

by Kluwer Academic Publishers, 

101 Philip Drive, Norwell, MA 02061, U.S.A. 

In all other countries, sold and distributed 

by Kluwer Academic Publishers Group, 

P.O. Box 322, 3300 AH Dordrecht, The Netherlands. 

Publication dates 

Part 1 20 Jan. 1984 
Part 2 21 May 1986 
Part 3 19 Oct. 1986 
Part 4 3 Aug. 1989 

All Rights Reserved 

© 1989 Kluwer Academic Publishers 

No part of the material protected by this copyright notice may be reproduced or utihzed in any form or by 

any means, electronic or mechanical, including photocopying, recording, or by any information storage and 

retrieval system, without written permission from the copyright owner. 

Printed in the Netherlands 



Title-page (3) 

Conienis (5) 

Dedication by C. G.G.J, van Steenis (7) 

Abbreviations and signs (41) 

in alphabetical sequence 

Alseuosmiaceae by C. G.G.J, van Steenis 335 

Araucariaceae by D.J. de Laubenfels 419 

Aristolochiaceae by Ding Hou 53 

Chloranthaceae by B. Verdcourt 123 

Chrysobalanaceae by G.T. Prance 635 

Coniferales by D.J. de Laubenfels 337 

Cruciferae by B. Jonsell 541 

Ctenolophonaceae by A.M.N, van Hooren & H.P. Nooteboom 629 

Cupressaceae by D.J. de Laubenfels 443 

Elaeagnaceae by J.F. Veldkamp 151 

Ixonanthaceae by R. Kool 621 

Linaceae by A.M.N, van Hooren & H.P. Nooteboom 607 

Magnoliaceae by H.P. Nooteboom 561 

Menispermaceae by L.L. Forman 157 

Monimiaceae by W.R. Philipson 255 

Olacaceae by H . Sleumer 1 

Opiliaceae by P. Hiepico 31 

Pinaceae by D.J. de Laubenfels 447 

Podocarpaceae by D.J. de Laubenfels 351 

Polygalaceae by R. van der Meijden 455 

Sabiaceae by C.F. van Beusel^om & Th.P.M. van de Water 679 

Sphenostemonaceae by C. G.G.J, van Steenis 145 

Taxaceae by D. J . de Laubenfels 347 

Trimeniaceae by W.R. Philipson 327 

Triuridaceae by J. P.M. van Meerendonk 109 

to volumes 4-10 

Addenda, corrigenda et emendanda by C. G.G.J, van Steenis el al 717 

fndex to scientific plant names by E.E. van Nieuwkoop 721 



l^r\ 11 1.; jj (,. ..: liVUU Y lbiJ\J JJ -L ii l;l-cj^ 

Dedicated to the memory of 


Many botanists must have wondered why as yet no volume of Flora Malesiana was dedicated to 
the outstanding botanist Carl Ludwig Blume, undisputed pioneer in planning the compilation 
of a 'Flora Malesiana'. 

The writing of this Dedication would have been greatly facilitated if a full biography of Blume 
had been existent, but none is available; there is not even a bibliography of his works. Only recent- 
ly, in 1979, two biographical attempts were made, by J. Maclean and by A. den Ouden, but 
only for the period 1820-1832; together with other biographical and obituary notes they are here 
assembled in Appendix B. I have also compiled a bibliography: Appendix A.* 

There are various reasons to account for the lack of data. At Leiden there are, in the Rijksher- 
barium archives, only few letters addressed to or written by Blume, and this is also the case for 
the University archives. Also Treub (B) in his papers on the history of the Botanic Gardens at 
Bogor complained about the lack of correspondence of Blume. The largest source of (official) 
letters is contained in the huge 'Rijksarchief at The Hague, but it will require a large, time- 
consuming effort to unearth these (D: 5). Blume's large private library was auctioned at Leipzig 
in March 1863, soon following his death, by the firm of O.T. W'eigel (B; D: 9). 

It has sometimes been suggested that the lack of a full biography - to which Blume was certain- 
ly entitled - could be explained by the fact that Blume had few friends (D: 8) and that his contem- 
porary colleagues were antagonistic. But this explanation does not really hold, as a biography of 
the charismatic Miquel was not written before a century after his death. In the Netherlands the 
climate is not favourable for biographies of scientists, at least not in botany (D: 7). 

For the reasons given above I have waited a long time to frame a worthy dedication, in the hope 
that some historiographer would feel attracted to compose a full biography of In the 
absence of this I have ventured to accumulate material myself, recently supported by a study of 
Maclean (B) on Blume's years in Java and based on archival research in the 'Rijksarchief, and 
an unpublished essay by den Ouden (B) on the same period based on details from several hun- 
dreds of letters in the same archives. 

To my regret biographers frequently do not give sufficient attention to personality and motiva- 
tions, but confine themselves to an appreciation of achievements. I have tried to form an opinion 
about this facet of Blume. From Blume's profuse writings much can be learned about his motiva- 
tions and his attitude towards society and people. It stands beyond doubt - and that must soon 
have been realized by his contemporaries (E, F) - that in the science of botanical taxonomy Blume 
was on a level with the great taxonomists of the previous century. But in the eyes of his close col- 
leagues he was an autocratic, dominant, unsympathetic person, and this impression still lingers 
around his name and overshadows the singular value of his scientific work. His sharp pen and 
especially his fanatical pursuit of a monopolistic position for the Rijksherbarium estranged him 
from his surroundings. Goddijn (B: 1931) has pointed this out very well. 

.My purpose in composing this dedication is to give a sketch of Blume's life, his work and his 
motivations in a detached way. Blume has a right to an impartial judgement; activities and per- 
sonalities should be kept well apart. In a few cases, where there is lack of clarity about the inter- 
pretation of historical data, I will give Blume the benefit of the doubt. 

(1) Shortly before his death in May 1976 the author of this Dedication and former Editor of Flora Male- 
siana, Professor C.G.G.J. van STthsis, finished the text of the manuscript. He had the intention to use this 
biography of Blume to conclude volume 10 of the Flora. Wc wholeheartedly like to carry out his intention 
here. - The General Editor. 

(2) The documentation here presented is recorded in six appendices: A. Bii'mf's publications annotated; 
B. Biographical sources; C. References to cited literature; D. Notes (mostly additional information considered 
useful to illustrate the situations under which Blume had to work, his surroundings, personalia, etc.); E. 
Eponymy; F. Honorary distinctions and memberships. 

The photograph on the opposite page is copied from Rumphia } (1847), Bi.umi silting above his treasures 
of the Javanese flora, including Nepenthes, Rafflesta, Rhizanthes, orchids and a rattan, presumably Plec- 
lotomia, the picture dating from the lime when he was at the height of his career. 


Flora Malesiana 

Towards the end of the 18th century two earUer attempts to compile a Flora of Malesia were 
made, namely by Francisco NoRONa in 1786 and by Louis Deschamps in 1794-1798 (B: van 
Steenis & VAN Steenis-Kruseman, 1970; C: van Steenis c.s. , 1954). Both attempts were abortive 
by the unfortunate loss of the material these collectors had made. 

In the early 19th century the time had come for the more thorough exploration of the tropical 
floras, both in the New and in the Old World. In the Indies it was started by W. Roxburgh and 
N. Wallich. In Malesia there had been quite some botanical activity in Raffles' time, notably 
by W. Jack in Sumatra and by Th. Horsfield and L.T. Leschenault de la Tour in Java, but 
these researches had led only to publications by Jack. 

The proper achievement fell to Blume, after the establishment of the Botanic Gardens at Bui- 
tenzorg (Bogor) in 1817, where a year later Blume started a research period of seven years which 
led to his brilliant scientific career. 

Carl Ludwig Blume, born at Brunswick (Germany) on 9 June 1796, was a son of the merchant 
Christiaan Nicolaas Ludwig Blume and of Melusine Caroline Sophie Drechsler. His father 
died before he was born and his mother died when he was five years old. He was an eager boy 
and was attracted by the study of pharmacy. To a high degree he was interested in travel books 
of foreign countries, a trend and interest possibly strongly developed in Germany since Hum- 
boldt's time, known as the 'Wanderlust', a tendency perpetuated to the present day (D: 10). 
Blume's interests were probably directed towards the many unexplored regions of the globe, in- 
cluding the tropics. By 1813 he used his heritance to buy clothes and equipment, and enlisted as 
a volunteer in the 'Liitzowsche Jagercorps', fighting against the French. Later on he went to the 
Netherlands, where on 29 December 1814 the Medical Board of the Dutch Forces appointed him 
as a military apothecary of the second class. On 6 April 1815 he was placed with the ambulance 
of the second division of the mobile forces in Belgium. He was present at Waterloo. According 
to the military Stamboeken (Registers) he was an apothecary of the second class in the hospitals 
at Den Helder and Leiden between 1814 and 1817. 

When in 1815 Prof. S.J. Brugmans was commissioned to bring back the collections of natural 
history from Paris to the Netherlands - collections which the French had taken there in 1795 - 
Blume was appointed as his assistent. 

In some way or other, young Blume enjoyed the support of the Duchess of Braunschweig, 
financially and otherwise. She fostered his career and had recommended him to Prof. Brugmans 
(t 1819), who urged Blume - who had performed his task excellently - to study natural history 
and medicine. Blume followed this advice and took a degree as Doctor of Medicine on 9 July 1817 
at Leiden (A: 1817). Shortly before this date, apparently in view of his doctorate, Blume finished 
his activities as an apothecary in the hospital at Leiden. On 17 October 1817 he returned in the 
service of the hospital as an M.D., after having obtained, on 6 October 1817, the degree of a 
health-officer of the second class of the forces and hospitals. On 1 1 January 1818 he was honour- 
ably discharged as a surgeon-major and on 28 March 1818 became a health-officer of the second 
class of the forces in the Netherlands East Indies. On 28 May 1818 followed the same appointment 
for the first class; he worked at Leiden till 17 March 1818. 

Shortly after his arrival in Java, on 1 1 January 1819, Blume was appointed deputy-director un- 
der C.G.C. Reinwardt in charge of the organization of Education, Medical service, Agriculture, 
Arts and Scientific investigation. He was then only 22 years old, but obviously highly esteemed 
for his ambition, zeal, knowledge and energy. His initial salary was / 500 annually. He lived in 
Reinwardt's house at Buitenzorg (Bogor), enlarged for this purpose, in the Botanic Gardens. 
He married the rich Wilhelmina Nicolasina Cranssen. This marriage was obviously not very 
successful. He was divorced in April 1830 in Brussels and he remarried at the end of that month 
Johanna Alletta Wilhelmina Waardenburg, by whom he had 7 children. 

At that time the Government was much concerned about serious tropical diseases, small-pox, 
typhoid, cholera, and in 1820 Reinwardt wrote a detailed report on the state of vaccination in 



the years 1818- 1819. All civil servants were informed of the Government's intention to maintain 
and promote vaccination. Blume was provisionally appointed 'Inspector of Vaccine' in 1819. He 
informed the Government that it was desirable to use indigenous plants instead of imported 
medicines which often lost their value during the long sea-voyage, and the Government requested 
him to make proposals. 

In the seven years between 1819 and 1826 Blume travelled widely in West and Central Java, 
as far east as Rembang, often accompanied by assistants, draughtsmen and interested persons, 
collecting plants and also animals; gathering information on all sorts of aspects, including the 
medicinal value of certain plants, inspecting epidemics, etc. ; in short he was engaged in an overall, 
thorough scientific exploration. 

He gathered many duplicates and his herbarium specimens are still in excellent condition. It has 
never become clear to me how these early explorers managed to dry and preserve their collections 
so well in the everwet tropics under the primitive conditions of the time, trekking from camp to 
camp (B: van Steenis-Kruseman, 1950). 

In 1821/22 he was in Bantam in the company of the civil servant J.B. Spanoghe; in 1822 he 
made a large exploration of Mt Salak ; in 1 823 of Mt Gedeh; in 1 824 he made a large tour of inspec- 
tion in the company of the clerk G.H. Nagel, the gardener W. Kent, and the draughtsman A. 
Latour, to many places: Kuripan (near Bogor) with hotwells in limestone then surrounded by 
primary forest, Mt Seribu (hills SW. of Jakarta), then to the Krawang region (E. of Jakarta) 
eastwards to Indramayu, proceeding to Cheribon, ascending Mts Tjeremai, Tangkuban Prahu, 
Burangrang, going as far as Tegal. Furthermore, he explored the then completely forest-clad large 
island of Nusa Kambangan (S. Central Java) where he detected Rafflesia. In 1825 he was again 
in Rembang (Central Java), but also in Bantam, ascending Mt Parang. 

These must have been hectic, creative years in Blume's life. In view of later controversies I have 
listed these explorations, which show that Blume covered a considerable part of West and Central 
Java, and that his travels partly covered the same habitats which had been visited by Kuhl and 
VAN Hasselt, members of the 'Natuurkundige Commissie', but also went beyond these. The 
result was of course that the majority of species were collected by both parties. 

In all probability Blume studied, analyzed, and described his collections in situ, facilitating 
later publication. In addition to all this field work he published scientific reports on many of these 
explorations, in part made public in a number of letters which he wrote to the brothers Nees von 
EsENBECK, published in the Regensburg journal Flora (A: 1823-1826). 

Finally he compiled all this material in the voluminous Bijdragen (A: 1825-1827), containing 
the concise treatment of some 700 genera and about 2400 species, belonging to 170 families of 
flowering plants. This achievement is colossal, as he had only very few books at his disposal, viz. 
Willdenow, Species Plantarum, Persoon, Synopsis, Sprengel, Anleitung and Systema Vegeta- 
bilium, DE Jussieu, Families des Plantes, Roxburgh, Flora Indica vol. 1 , and W. Jack, Malayan 
Miscellanies. He had of course also at his disposal the works of Rheede and Rumphius but they 
were of hardly any taxonomical use. He mentioned in his Enumeratio that he had seen the plates 
of Norona, obviously of a set since lost, but could not have had much profit from them for his 

The writing of the Bijdragen itself was a tremendous task, let alone the research incorporated 
in them, a great deal of the genera and species being new to science. This research work has ap- 
peared to be of very high quality, testified by the fact that a very large amount of his newly pro- 
posed genera still stand and that others, now merged with earlier described ones, were always good 
taxa and were later often still recognized as subgenera or sections. A great merit was that Blume 
hardly ever failed to recognize their proper affinity and almost always placed them in the proper 
family, evidence of his great systematic capacity. In view of the rather primitive state of tropical 
botany in his time this deserves great respect. Blume's skills in this field also appeared from a first 
attempt to construct a system of affinity for tropical orchids, laid down in the Tabellen en Platen 
voor de Javaansche Orchideen (A: 1 825), issued in part simultaneously with fascicle 6 of the Bij- 


Flora Malesiana 

dragen. He complained that he had had no access to contemporary literature on the family by 
R. Brown, C.S. Kunth, and L.M.A. du Petit Thouars, which he only received during the print- 
ing of his own system of the orchids. This first attempt was much later crowned with his 
monograph Flora Javae. Nova Series (A: 1858-1859) of the Orchidaceae, the largest and least 
known family of the Malesian flora. 

In addition he published in the first five fascicles of the Bijdragen data on the useful and 
medicinal plants of the families treated. 

Apart from his work with vaccination and his exploration and botanical research work, another 
duty had fallen to Blume, when he was in June 1822 appointed director of the Botanic Gardens 
at Buitenzorg (Bogor), at Reinwardt's request succeeding him. Reinwardt himself repatriated 
in that year. The annual salary was/ 1000. This was a task in itself; besides enriching the garden 
with plants he collected during his own travels, he was also in contact with other gardens abroad, 
for instance at Mauritius and Calcutta, with the purpose of exchange. 

Blume was well aware of the fact that he should attempt to stimulate our consulates in foreign 
countries to collect plants or seeds for the garden, a policy which he later also followed when he 
was director of the Rijksherbarium. For the Buitenzorg garden he wished to have more Chinese 
and Japanese species and to obtain material he wrote to the Dutch consul in Canton and the 
representative in Deshima (Japan). A year later, in 1824, he instructed the Dutch in Japan how 
to dispatch seeds and plants to Batavia (Jakarta). 

Furthermore, Blume sent Javanese and other exotic plants in small baskets to the university 
gardens at Leiden, Utrecht and Ghent in the Netherlands and also dispatched seeds to the 'Societe 
de Flore' at Brussels. 

In 1823 Blume published the first Catalogus van ... 's-Lands Plantentuin te Buitenzorg (A). 
In the listing were many manuscript names of Reinwardt under the latter's name. Blume himself 
added several new genera under his own name with valid descriptions. Without doubt Blume was 
the botanical 'motor' of this catalogue, Reinwardt having been too much occupied by ad- 
ministrative and organizational matters, and besides having been previously occupied by his large 
exploration of eastern Malesia. It should be added that Reinwardt's plant-systematical 
knowledge was meagre (D: 8). 

On 11 June 1822 Blume was also definitively appointed as 'Inspector of Vaccine' and had to 
attend to his medical-pharmaceutical duties as well. He reported on the virtues of hotwells in 
Krawang (A: 1825, 1839), gained information on the fight against cholera, etc. for which he in- 
itiated medical treatment, and paid attention to medicinal plants (A: 1825, 1832). On 12 August 
1823 he was appointed commissioner of the civil health service. In short, his duties were manyfold 
and his achievements in these years are of tremendous proportion. 

In 1824 Blume received permission to extend his research to all Dutch possessions in the East 
Indies and was allowed to publish in the journals of Dutch societies. The Government would pay 
for the printing of a book on botany, obviously the Bijdragen, with the provision that all 
discoveries, observations and prepared specimens would be the exclusive property of the Nether- 
lands Government. 

In a letter dated 6 August 1825, no. 365, Blume informed the Governor-General about the pro- 
posed publication of his large book Flora Javae, pointing out that this was urgent as other persons 
who had explored in the Netherlands Indies were already active in having their discoveries printed. 
These other persons were obviously the French explorer L.T. Leschenault, the American Th. 
Horsfield, and especially the British W. Roxburgh and W. Jack. He said that with the insecure 
life in the tropics, when so many fell an early victim to tropical diseases, he felt that he had to 
safeguard his research, the result of his extensive field work and observations for science. 
Therefore he had decided to publish the very concise Bijdragen in anticipation of the large work 
Flora Javae which he had in mind. He mentioned that his own health slightly deteriorated, but 
there is no evidence that he was ever seriously ill in Java (D: 11). The Bijdragen were certainly 
not merely a striving for priority. 



It was then that years of negotiation started about financing the expensive Flora Javae. For its 
elaboration he requested a leave of three to four years in Europe, necessary for the acquisiton of 
information which the new literature and the comment of experienced botanists could offer, and 
this required visits to some of the famous herbaria in Europe. He offered to stay in Europe on 
part of his salary. 

In September 1825 the Governor-General permitted him a two-years stay in the Netherlands, 
at one third of his pay. After a frustrated attempt of Blume to ship a large amount of living 
material to the Netherlands, and an offer to pay for his own passage, the Government finally 
decided by 26 June 1826 to commission Blume for two years leave to the Netherlands on half-pay. 
His medical activities and the vaccination were assigned to his colleague Peitch and the botanical 
work in the Gardens would be looked after by the gardeners James Hooper and Alexander Zip- 
pelius who, together, would be paid from the other half of Blume's salary. These were times of 
poor economy in the kingdom. 

Blume took with him 29 cases of herbarium material, sailing in the ship 'Christina Bernardina', 
destination Brussels, then the capital of the kingdom. He had the good fortune that the ship ar- 
rived safely, so many earlier dispatches having been lost by shipwreck, for instance several of 
Reinwardt's. By the end of 1826 Blume arrived in Holland. By far the main part of the collec- 
tions were made by himself, minor ones were included, e.g. those made by Reinwardt in Java 
and East Malesia (Celebes, Moluccas, Timor), local Javanese collections made by the gardeners 
ZiPPELius, Kent and Hooper m the vicinity of Buitenzorg, etc. It should be stressed, however, 
that none of the collections of Kuhl and van Hasselt were included, as these were property of 
the 'Natuurkundige Commissie'. Later, in 1828, these latter collections were dispatched to the 
Museum of Natural History at Leiden by G. van Raalten, who had been taxidermist in the ser- 
vice of the 'Natuurkundige Commissie', assisting Kuhl and van Hasselt. Van Raalten was also 
a capable draughtsman; he died at Kupang (Timor) in 1829. 

Van Breda's archive, now at the 'Hollandsche Maatschappij', Haarlem, contains a partial 
abstract of a letter dated 22 July 1825 by G. van Raalten (B: 1825), in which he complains that 
Blume - who had inspected the orchids in the Kuhl & van Hasselt herbarium - had noted which 
species had been depicted of their collections. He became afraid that Blume's pubUcation would 
precede the publication of the Kuhl & van Hasselt plants and found this unfair. He felt extreme- 
ly sorry for the misfortunes which befell Kuhl and van Hasselt. This letter was certainly one 
of the arguments for later, unjust accusations that Blume stole scientific property. Van Raalten 
pointed out that Blume had agreed with van Hasselt to work out the orchids jointly, which 
Blume also acknowledged in his Bijdragen; in fact some 27 names have a dual authorship, as I 
have elucidated (B: van Steenis, 1980). As a non-botanist van Raalten did not understand that 
in such unfortunate situations the dead have no claim unless they left manuscripts. 

A testimony that Blume, after his departure from Java, had no access to manuscripts or draw- 
ings of Kuhl and van Hasselt is the fact that in the Bogor Library there is - or at least was, 
before World War II - a book containing drawings of Kuhl & van Hasselt (on Asclepiadaceae, 
Orchidaceae, etc.); it is a further proof that Blume did not have these documents (D: 1). 

Still, the letter by van Raalten, which was badly understood and interpreted, had influence. 
Accusations and slander lead a long life, and are often eagerly reproduced by antagonists. Thus 
even Temminck, the director of the Museum of Natural History at Leiden, wrote in 1 828 - when 
the Kuhl & van Ha.sselt herbarium was transferred to Blume - that the latter should guarantee 
priority to the manuscript names of Kuhl & van Hasselt in publishing, although Temminck must 
have been quite well informed about the situation. I regret that Smit (B: 1979) in his essay still 
accepted van Raalthn's accusation. 

On the arrival of Blume in Brussels, he reported to I). J. van Hwijck (1786- 1855), administra- 
tor of Education, Arts and Sciences in the Uepartmcnt of the Interior, who was very much im- 
pressed by Blume's personality and works. In December 1826 van Ewijck spoke highly of 
Blume. praised his diligence and knowledge and declared himself in favour of the /'lora .lavae 


Flora Malesiana 

plans. The Minister contacted his colleague of the Colonies, who in his turn applied to King 
Willem I. This was followed by endless discussions who would pay for the publication of Flora 
Javae. The result was that Blume received 7000 florins and that the Dutch Government would 
buy 50 copies (5 florins for each instalment), the Netherlands Indies' Government would buy 4 
copies, and that he was allowed to appoint a draughtsman (Arckenhausen) for a period of four 
years. Blume had in mind to publish 250 instalments. 

In the meantime Blume pursued his activities in Holland, continued the Bijdragen, and com- 
posed a new work under the title Enumeratioplantarum Javae . . . (A: 1827-1828). The treatment 
was more elaborate than that of the Bijdragen. It was published in Leiden. He mentioned on the 
title page that he had also used material from Kuhl and van Hasselt, but this is hardly possible 
as this came only available to him in 1828 (D: 1). 

Blume dedicated the first volume to the Nees von Esenbecks at Regensburg, with whom he 
had early friendly relations for several years. Blume's frequent letters to them on his experiences 
in the exploration of Java were published in several volumes of the journal Flora, and he frequent- 
ly sent them cryptogams, mosses and fungi; when he returned to Holland in 1826 he stuffed empty 
spaces in his cases between his parcels with moss samples, especially hepatics, which enabled 
Th.F.L. Nees von Esenbeck to publish on Javanese Hepaticae in 1830. Partly out of courtesy 
the latter published a paper on Javanese Fungi, with Blume as co-author (A: 1827). As a matter 
of fact Blume extended his interest distinctly to cryptogams, and earlier had already pictured and 
studied mosses and fungi himself in the field. This interest did not wane, because in 1841 he readily 
agreed with Zollinger to buy lichen collections from Java where Zollinger intended to explore. 

The second volume of the Enumeratio, dedicated to W.J. Hooker, consists mostly of descrip- 
tions of Pteridophyta; in fact it is the first account of them in Java. It proves Blume's thorough 
botanical knowledge, because he was mostly versed in Spermatophyta. Notwithstanding that, this 
volume is as complete and its contents as accurate as that of the flowering plants, according to 
Hennipman (C: 1979). 

When in 1828 Blume's leave came to an end, he requested discharge of his position as chief 
of the Civil Health Service. This was granted because he would continue to work on botany and 
would not return to Java. 

By Royal Decree of 22 June 1828 he was granted from 1 July 1828 onwards an annual salary 
of 3000 florins for his services and an annual half-pay of 2000 florins, till he had obtained 
another position. Blume had to cede in this same year his immense collection of animals and 
insects to the Museum of Natural History at Leiden. As compensation he would receive an an- 
nuity (B: GiJZEN, 1938). 

The first two parts oi Flora Javae appeared in Brussels, in 1828, under authority of Blume and 
his adjunct. Dr. J.B. Fischer. J.G.S. van Breda (C: 1827-1829), then professor at Ghent and 
by profession a zoologist, would participate, or at least elaborate, the Orchidaceae and Asclepia- 
daceae. For this purpose the drawings and descriptions of plants made by Kuhl and van Hasselt 
were also given to van Breda. 

On 31 March 1829 the Rijksherbarium was founded at Brussels, with Blume as director, with 
the title of professor. One of his first actions was to instigate that the Botanic Gardens at Buiten- 
zorg should regularly provide consignments of plants to the Rijksherbarium, and furthermore, 
that the members of the 'Natuurkundige Commissie' in the Indies should not distribute specimens 
to foreign herbaria. 

The Rijksherbarium did not long exist at Brussels because of the 1830 rebellion, and was saved 
in the nick of time and transported to Leiden by Fischer and von Siebold. This subject has been 
fully reported by my wife (C: van Steenis-Kruseman, 1962). Blume himself was not on the spot, 
because he was on his honeymoon to Geneva. He combined this tour with the object of inspecting 
the Roemer herbarium, which was for sale, to see whether it was worth-while to purchase it for 
the Rijksherbarium collections. 

The Rijksherbarium, after its transfer to Leiden, was at that time not affiliated to the Universi- 



ty, but was subjected immediately to the Ministry of the Interior. That Ministry drafted an In- 
struction for the director, effective from the first of January, 1831 (C: van Dam, 1832). 

Blume continued the issue oi Flora Javae. Mid- 1830 35 instalments had been published. Unfor- 
tunately, the subscriptions appeared insufficient and money ran out, and the work was temporari- 
ly abandoned. 

Blume did his best to expand the Rijksherbarium collections on a large scale. Via the Ministry 
of Foreign Affairs he urged civil servants abroad and in the colonies to collect plants and make 
herbaria. ' For this purpose he composed a booklet of instruction (of which I have not been able 
to trace a copy) on how plants should be made into a herbarium, as drying plants in the tropics 
brings along difficulties by the moist climate and the often bulky and/or fleshy structure of the 
material. Moreover, there was the problem of frequent insect damage once plants are dried. 

With some people Blume succeeded. There is e.g. a large collection of several hundreds of 
specimens made by the Dutch consul in Venezuela, J.G. van Landsberge, made in 1842. This 
collection is arranged by families, but remains unidentified to the present; it contains many 
duplicates. On the whole, however, Blume's urging did not meet with great success. 

Blume also approached missionaries to collect plants in their territory, and stimulated phar- 
macists to do the same; those whom he tutored at Leiden he gave special attention and instruction. 
Although in this way many people sent overseas were aware of his wishes, the results were very 
meagre, as compared for example with the results of F. von Mueller in Australia in his contacts 
with missionaries. The latter's success is probably to be ascribed to the fact that he maintained 
a very regular correspondence with them and kept them timely informed of results. Besides, von 
Mueller lived much closer to them. 

In general, the attempt to acquire botanical material by stimulating an interest in the tropical 
flora among medical men and other residents in the colony and the collecting of specimens was, 
as far as I can judge, not successful either. The endeavour in itself was excellent, but possibly 
precocious in the early 19th century. 

In addition Blume was engaged in buying collections which were for sale. A curious, significant 
example was a collection of Javanese plants offered in 1837 for sale to the Government by the 
German physician J.G.H. Kollmann, who was in the service of the Dutch East Indian army. This 
offer was referred to Blume who found to his great surprise that this collection contained also 
the set of duplicates (about 4000 specimens) which he had conscientiously left at the Botanic 
Gardens in Buitenzorg (C: van Steenis-Kruseman, 1950; D: 4). 

It should be borne in mind that it was factually impossible for Blume to work on incoming col- 
lections without having a large staff of botanists at his disposal. From numerous letters in the 
'Rijksarchief it is evident that he pleaded time and again for the appointment of staff officers. 
Notwithstanding the esteem he was held in by the Ministry of the Interior and the sympathy of 
some high officials, notably van Ewijck, it was of no avail. He could not even attain a permanent 
position for his two closest collaborators. Dr. J.B. Fischer and his draughtsman and handy-man 
J.C.P. Arckenhausen(C:Griepc.5., 1977; D: 12). Financially the Netherlands were at that time 
at low ebb. Blume, moreover, was unfortunate with respect to the few scientific co-operators he 
had. Van Hasselt and Flscher met untimely deaths and van Breda took another job. 

Members of the 'Natuurkundige Commissie' were entitled to work out the results after seven 
years of exploration in the East Indies, but in this category it was only P.W. Korthals who per- 
formed excellent work. Korthals was possibly a modest man, in the shadow of Blume, but his 
work in the field and in science was of the same high quality. Korthals would have been an ex- 
cellent staff member, but after his retirement he devoted his time to philosophical contemplation. 

(1) A. DE Candollf. mentioned (B: 1862) thai Biomi; told him the Netherlands Indies' Government had 
ordered, at Blumk's request, thai all physicians in their service should have A. P. di ("ani)oi.ij:'s essay Sur 
les propnitis des planles (1816) as a botanical guide. 


Flora Malesiana 

as became evident from Zollinger's diary. Blume cannot be blamed for the fact that Korthals 
abandoned botany (B: Zollinger, 1841; D: 8). 

H. VAN Hall had worked with Blume on a temporary basis from about 1850, but was only of- 
ficially appointed adjunct-director in 1854, the only permanent scientific collaborator Blume was 
ever allowed. 

Deficiency of technical staff was another drawback; here again attempts to expand failed. 
Apart from his draughtsman Arckenhausen technical assistants were few. This must in my opin- 
ion be one of the main reasons that hardly any duplicates were distributed in order to exchange 
material with foreign institutes to enrich the Rijksherbarium. Foreign colleagues complained that 
Blume asked for their material, but seldom gave a return. This greedy and monopolistic attitude 
made him unsympathetic. Evil tongues claimed that it was Blume's intention not to distribute 
duplicates, as he wanted to prevent new species to be described by others. I cannot believe this 
to be correct for in any case he could have distributed duplicates of species already described by 
himself. Obviously Blume was not in favour of seeing undescribed species published on dupli- 
cates. Not until the 1860s, under Miquel's directorship, numerous duplicates were distributed, 
partly unnamed. The same open policy was followed by the Herbarium Bogoriense (with an ex- 
ception of selected Javanese collections made by C.A. Backer) and this more generous attitude 
was also kept up by Merrill in Manila, and from Miquel onwards by the Rijksherbarium. In 
the first place this is done for the greater safety of the collections (in that respect we have but to 
think of the disastrous effect of the fires in Berlin and Manila) but also because all research on 
Malesian plants must be welcomed, irrespective where and by whom. It is self-evident that in case 
of free-for-all descriptions a lot depends on the quality of the collaborators. It is true that not 
infrequently mediocre or uncritical collaborators have created more extra work rather than solved 
the problems for their successors. 

A great inconvenience associated with duplicates of the early Dutch collectors was the fact that 
they were not numbered, neither by Blume himself, nor by Korthals, Reinwardt, or others. 
Through this, typification is difficult and it is sometimes impossible to know which duplicates 
belong to which collection. The more praiseworthy a Teijsmann, who consecutively numbered the 
Buitenzorg collections! But then the latter had more personnel. Blume's limited staff was certain- 
ly one of the reasons that the numerous collections remained undivided. Whom could he trust to 
distribute the unmounted collections in a responsible way? 

As already mentioned it was not before the 1860's that Miquel, Blume's successor, instigated 
the policy of free distribution of duplicates, but certainly did not do it himself; he had it done 
by technical personnel. Without doubt the distribution of duplicates was extremely important as 
the result was the acquisition of numerous duplicates in exchange from foreign herbaria in 
Europe. It is a pity that this distribution lacked carefulness as regards the labels. Occasionally 
specimens with Blume's handwriting were sent elsewhere, for instance to Paris, while specimens 
retained at Leiden have labels written by a clerk. This is not seldom a nuisance in connection with 
the assignment of the holotype. Sometimes the use of wrongly printed labels is confusing, for in- 
stance of Korthals specimens of which sometimes 'Sumatra'-labelled plants are really from SE. 
Borneo. At Blume's and Miquel's times most of the Malesian collections were not mounted; this 
was only done towards the end of the 19th century. 

As the prime botanist he was, Blume's interests were by no means restricted to those of a scien- 
tist working in seclusion. He was always keen on the development of the colony towards better 
living and status. He stressed the importance of promoting the cultivation of plants not only in 
the interest of big enterprise, but he held the opinion that there had to be a balanced situation 
for the benefit of all! This comprised also the introduction of new, useful plants. If one reads his 
general papers it appears that he had wide interests from his early stay in Java onwards. In the 
first five instalments of the Bijdragen he provided families with notes on their useful species. He 
wrote a monograph on the peppers (A: 1826) and as early as 1820 he took the initiative to advise 
the Government on the importance of cultivating indigo and of importing cochineal, and last but 



not least to import Cinchona, which materialized only three decades later through Hasskarl. In 
many papers he advocated more activity in agricultural matters and stressed the importance for 
national well-being in commercial, hence financial, aspects, for the Dutch as well as for the native 

As a medical man, in his capacity of 'Inspector of Vaccine' , and during his many travels, Blume 
was of course in intimate contact with the Javanese people and he took their welfare as much to 
heart as that of the Dutch people; he clearly regarded them all as co-citizens. For example, he 
pleaded openly in a letter to the Governor-General of the Indies (A: 1829) for the desirabihty of 
abolishing opium, as he found this a menace for the population; only much later this was 
regulated indeed by the Opium Law. 

In 1842 Blume founded, together with Ph.F. von Siebold and on the instigation of J. Pierot, 
the 'Societe Royale pour I'Encouragement de I'Horticulture dans les Pays-Bas' (Royal Dutch 
Society for the Advancement of Horticulture). This was part of his endeavour to make botany 
subservient to the general interest of the kingdom and to create a stimulant for new financial and 
commercial interests. In a first issue of the above-mentioned 'Societe' (A: 1844) he compiled a 
large list of useful plant species. Also later he showed his unfailing devotion by a stimulating paper 
on timbers resistant to pile-worm (A: 1859). 

Altogether he held enlightened, progressive ideas - not so popular in those days - and in his 
opinion the native people ought to have their share of welfare, not in the least because their man- 
power was an essential aspect of a prosperous colony. In this respect it is significant that he named 
the genus Santiria after Bapa Santir, an old Sundanese, who accompanied Blume on his explora- 
tions of Mt Salak. It was Junghuhn who took this amiss (B: Junghuhn, 1853) and suggested that 
Blume was consciously deceptive in pretending to be generous, but really threw a blame on great 
botanists and other dignified man who were the only persons entitled to be honoured by eponymy. 
In his colonial arrogance Junghuhn called Bapa Santir an inferior person, not more than a sim- 
ple 'pakkedrager' (kuli, carrier), whereas in all probability Bapa Santir was an intelligent man 
and an outstanding local authority on plants who knew his way in the forest, knew the vernacular 
names and uses of forest plants and assisted Blume in many ways. It is testimony of the irony 
of fate because in history Junghuhn is reputed to be the pioneer and advocate of a progressive 
society of freethinkers, whereas Blume is remembered as a distinctly conservative person, though 
all his writings give evidence of a progressive, liberal mentality. It appeared that Blume, mirabile 
dictu, was the more enlightened of the two; he was certainly devoid of any racial prejudice. 

In 1843 Blume started the journal De Indische Bij, another endeavour to promote an interest 
among the Dutch public in the understanding of the colony. Only one volume was issued (1843), 
mainly filled with papers by himself and his friend C.F.E. Praetorius, Director of Cultures in 
Java, on all kind of subjects, partly political, partly ethnographical, on Borneo and South 
Sumatra, and on plant fibres. 

Returning to Blume's scientific works: in spite of the untimely abandoning of his Flora Javae, 
he set up another large-scale work in the thirties, Rumphia, the scope of which included also other 
parts of Malesia. The first fascicle appeared in 1836. It consisted finally of four volumes 
(1836-1849). This work was of the same critical standing as Flora Javae, to which it was similar 
in size and printing. In a sense it is an attempt towards a Flora Malesiana. Towards the end of 
the forties Blume again managed to issue some important parts of Flora Javae, namely the Filices 
(instalments 36-39 in 1847 and 40 in 1851) and the Loranihaceae (instalments 41 & 42 in 1851). 
How these issues and Rumphia were financed is unknown to me. 

The abrupt end of Flora Javae was regrettable and H.C van Hail, professor of botany at 
Groningcn, was much concerned about its continuation, which he found of national importance 
(C: van Hall. 1856). In a session of the Royal Academy of 28 June 1856 he proposed that this 
lofty body might form a committee to approach Blumi; in order to come to a proposal from the 
Academy to the Government for further financing Flora Javae; at that time 42 instalments, each 
with 6 plates, had been issued. I do not know if van Haii 's pleading led to any further action, 
but it shows that / lora Javae had supporters. 


Flora Malesiana 

After Blume's death there obviously remained illustrated printed material for a continuation 
of Flora Javae. These 23 coloured plates, called Planches inedites, mostly represented species of 
Loranthaceae and Ericaceae, all provided with analyses. They were offered for sale as a packet 
by the firm van der Hoek, Leiden, in 1862 or 1863 (A: 1863; C: van Steenis, 1947). 

Towards the end of the forties, when Blume was in his prime, he must have been disappointed 
with the untimely discontinuation of the two works on which he had set his heart. Flora Javae 
and Rumphia, and the insufficient public interest in his journal De Indische Bij. Moreover, clouds 
had gradually gathered round his claim that the Rijksherbarium had the monopoly for housing 
and possessing all collections made in the colonies by persons in the pay of the Government. He 
based this claim on the Instruction for the Rijksherbarium of 1832. This claim, however, was an 
optimistically exaggerated interpretation of art. 10 of this instruction which read (transl.): 'The 
Director will attempt to acquire collections, notes and drawings from all civil servants or people 
in the pay of the Government through proposals at the proper place and authority' (C: van Dam, 
1832). Blume may have had a moral right to claim these collections, but could not refer to a legal 
right. His claim was not attended to and this must have been a thorn in his flesh. 

Blume opposed the founding of Herbarium Bogoriense by Teijsmann in 1844, claiming that 
the latter should send the specimens to the Rijksherbarium, or at least the duplicates, but he found 
insufficient understanding with Teijsmann, who foresaw that he would have little profit from this 
in the way of a speedy naming of the specimens. Furthermore, Teijsmann's assistant, J.K. 
Hasskarl, had assembled a large private herbarium which he took with him on repatriation to 
Germany. Then von Siebold's herbarium was elaborated at Munich by Zuccarini (D: 2) where 
the types were left. W.H. de Vriese, professor at Amsterdam, had acquired the herbarium of 
Splitgerber, made in Surinam, but had not donated this to the Rijksherbarium. Finally, 
JuNGHUHN, officially belonging to the medical department in Java, had assembled a very large 
herbarium in Java, which Blume could not get into his hands (D: 3). It was purchased by Leiden 
University, under the condition that it should not be incorporated in the Rijksherbarium; it was 
entrusted to de Vriese. Finally, there was the rising star of tropical botany, F.A.W. Miquel, who 
originally published valuable monographs of Piper aceae, Cycadaceae, Casuarinaceae, Melocacti 
(partly for de Candolle's Prodromus), and later elaborated various large families in Martius' 
Flora Brasiliensis. He became also more and more interested in Asiatic plants, starting with his 
Analecta Botanica Indica, published by the Royal Academy. Miquel was a man of immense out- 
put and diligent handling of material, with an open mind for collaboration, which he brought in 
practice himself. Considering that, if the Junghuhn collection fell into Blume's hands, identifica- 
tions would be endlessly retarded, combined with Junghuhn's natural desire that it should be 
speedily worked out, de Vriese reasonably entrusted Junghuhn's collection for this purpose to 
Miquel. With elaborate support (e.g. Bentham's), the latter indeed published the Plantae 
Junghuhnianae. This must have caused immense irritation to Blume, who was constantly on the 
barricades defending his institute, stressing again and again that collections made by government 
officials with government money ought to be deposited in the Rijksherbarium. This monopoly 
also concerned himself. My wife (C: van Steenis-Kruseman, 1979: 51) wrote: 'whatever has been 
said to Blume's discredit, one thing is certain, and that is, that he was possibly the only botanist 
(and a devoted, not to say inspired one) in his period who had no private herbarium.' 

It is ironic but true that Blume's strict monopolistic claims made people reluctant to put their 
collections under his care, though Blume was, although not legally, at least morally in his rights. 
Even admitting that his claims were correct, it must be said that he should have realized that, if 
all these collections had been donated to the Rijksherbarium, he could as a single person never 
have mastered them. This would have been necessary, as some people wanted names and iden- 
tifications. He should have tried to compromise and initiate collaboration and division of labour, 
at least with Miquel and de Vriese, and not sit tight-fisted on propriety of collections. But ob- 
viously he could not well adjust himself to the changing conditions of the times and the rise of 
capable colleagues in his specialized field. This led to most unfortunate friction and a clash of 



personalities. He offended especially Junghihn in writing with his sharp pen an acid comment 
in Rumphia (1847 or 1849?) on Junghuhn's so-called Lycopodium arboreum which he had 'at 
first sight' recognized as belonging to the conifer genus Dacrydium, and Blume renamed Primula 
imperialis Jungh. as P. Kuhlii Blume, claiming that Kuhl had found this first and thus had priori- 
ty for eponymy, nomenclaturally wrong of course. Junghuhn complained that Blume begrudged 
him to describe Acer javanicum and had renamed this wrongly A. niveum, in which Junghuhn 
in turn was wrong. In short, about 1850 the fight was on and several very sharp and polemic 
papers were published to and fro (D: 13). 

The unfortunate result was that Blume became a still more isolated and probably a rather em- 
bittered person. Apart from odd fascicles of Flora Javae and Rumphia he had no opportunity 
for further great undertakings. He then put himself to proceed with a subject, stipulated in the 
1832 Instruction for the Rijksherbarium, namely compiling a catalogue of the collections of that 
herbarium. \s this implied identifications, this was not a clerical task for a non-botanist. My wife 
mentioned (C: van Steenis-Kruseman, 1979: 35) that the scientific arrangement of the collections 
was started by J. Pierot (1831-1840), who was succeeded by J.H. Molkenboer (assisted by C. 
Kerbert and Schultes Jr, the son of J. A. Schultes) (1840-1846), and finally by H. van Hall 
(1853-1862). Their work of course facilitated Blume's later Museum Botanicum. These helpers 
were named 'assistants', only van Hall was designated the title 'conservator'. 

In December 1850 Blume had to face an official new Instruction for managing the Rijksher- 
barium collections (C: Thorbecke, 1850). This was to meet official complaints by de Vriese, 
Junghuhn and von Siebold and especially Miquel, all influential persons, who wanted to bor- 
row material, requests only reluctantly given in to by Blume. Miquel wrote to von Schlechten- 
dal (B: Stafleu, 1970: 321): 'Es ist mir endlich gelungen, das Reichs Herbarium zu offnen. Nach 
einem Befehl der Regierung sind die Samml. aus Borneo, die noch ganz unbearbeitet waren, mir 
zur Disposition gestellt.' Blume was ordered to proceed with the catalogue; no unicates were to 
be removed from the collections; furthermore, the director had to refrain from publishing 
discoveries by still living persons of the former 'NatuurkundigeCommissie', unless with their con- 
sent. It must have irritated him considerably that responsibility and authority were restricted. 

The catalogue, named Museum Botanicum, was printed in fascicles, all filling one sheet (16 
pages), apparently with the intention to publish the fascicles monthly. It consists of two volumes 
in which the fascicles of volume one are dated by the year and month. The first volume was dated 
from 1849 to 1852 and finished with an index. The second volume was started with a fascicle dated 
1852, but fascicles 2-8 are undated, fascicles 9-16 being dated November 1855 to June 1856. 
There was no index; this was later composed by myself and Chew Wee Lek (C: van Steenis & 
Chew Wee Lek, 1974). As Beumee (B: 1948) and Stafleu & Cowan (B: 1976) have pointed out 
there are discrepancies about the dates of publication and this induced the latter towards sug- 
gesting that Blume withheld literature from his colleagues (Miquel, Weddell, etc.) and that in 
other cases the possibility of antedating cannot be excluded. Miquel (B: 1856) severely criticized 
the doubtful datings of the fascicles. It is quite probable that not every fascicle was for sale at 
the published date, but sold in lots, and confusion remains. In the absence of well-founded data 
regarding the authority which paid for the publication, who arranged the sale, and whether one 
could subscribe, we must refrain from further comments (D: 6). Possibly Blume still had a 
manuscript for one other fascicle which is known as Metan}>es botaniques (A: 1855). Up till the 
present it was assumed not to have been effectively published. This is, however, wrong, as I have 
discussed earlier (B: van Steenfs, 1986). The pamphlet was privately published and donated by 
Blimi; to his close friends; at least two copies still exist. 

The Museum Botanicum is an important, critical work; it contains some attempts towards revi- 
sions and, though species and genera from all over the tropics were dealt with, the main text refers 
to the Malesian flora. We do not know why the Museum Botanicum was rather abruptly abandon- 
ed. It is not unthinkable that Blume wanted to unburden himself from his old love, the Or- 
chidaceae, and saw an opportunity to publish this masterly work which he had had constantly in 


Flora Malesiana 

mind since his early Buitenzorg years. This work had been interrupted several times, first when 
his collaboration with van Hasselt came to an untimely end by the latter's death, and later by 
the early leaving of van Breda. Now it was published as Flora Javae, Nova Series (A: 
1858-1859). There is also a French-titled edition, with a preface in French, but otherwise iden- 
tical. According to W.E.G. Seemann (B: 1859) Blume complained that the Government had not 
contributed to its financing; obviously Blume, who was a man of means, had taken the risk of 
financing it himself. Besides the excellent works of R. Brown, Lindley and Reichenbach on the 
Orchidaceae and the affinities within the family, also Blume's work is very important and of 
similar standing, and naturally of special importance for Malesian botany. 

Blume, naturalized as a Dutch citizen in 1851, died in Leiden, after a long, painful illness on 
February 3, 1862, at the age of 65. 

As said before, Blume is through his large oeuvre - including eight important and critical 
botanical works of high standard: the Catalogus, the monograph on Piperaceae, the Bijdragen, 
the Enumeratio, Flora Javae, Rumphia, Museum Botanicum, and Flora Javae, Nova Series - one 
of the great botanists of the former century. A ninth treatise, on cholera in Asia (A: 1831), is 

His creative output is imposing. He distinguished eight new families, to wit, Apostasiaceae 
(now mostly judged a subfamily among Orchidaceae), Burmanniaceae, Cardiopteridaceae, 
Dipterocarpacaeae, Hernandiaceae, Myricaceae, Sabiaceae, and Schisandraceae. In addition he 
described, from Malesia alone, some 300 new genera of which 160 are still used, and 140 are now 
in synonymy, either for reasons of nomenclature or for new systematical insights. However, they 
were all proper taxa and are still frequently recognized as infrageneric taxa, e.g. Tarrietia and 
Campanumoea. Furthermore, he described his genera and species almost always in the proper 
families cq. genera, testimony of his systematic vision. 

As to his scientific achievement, his talent was soon recognized, both in Holland and abroad 
and he was soon made a member of learned societies (F). As usual for members of the 'Leopol- 
dina', they should have a cognomen; Blume took for himself the well-chosen name Rumphius 

Many generic names (E) and very many species were named after him. We are pleased that the 
journal of the Rijksherbarium, Blumea, is named after him. 

As an explorer Blume was exemplary in multidisciplinary approach by making observations on 
the spot, having a draughtsman with him, interrogating the native people about the uses and ver- 
nacular names, collecting insects and other animals, and paying attention to soils, mineral wells, 
etc., and by timely reporting about his field research, a good habit which young explorers of the 
present day should take more to heart. Through his medical profession he made also observations 
about native diseases and tried to cope with these to relieve suffering of the people. 

All his endeavours in this field and also his many advices on agricultural and horticultural af- 
fairs were focussed on tying up scientific botany and practice for the benefit of society. As such 
he was the opposite of the scholar in the ivory tower. His sharp observation power paired with 
interest were not confined to botany, as appears from his conclusions on serious contagious 
diseases among which cholera and typhoid were the most dangerous. As 'Inspector of Vaccine' 
he went to Central Java on inspection during a cholera epidemic and observed that the disease 
was especially prevalent in the lower lands, and less so in villages in the mountains. He deduced 
that cholera was spread by the polluted water and that the freshwater wells in the mountains were 
less contaminated. He prescribed all sorts of simple means for a diet and medicinal substances 
from native plants, but in the first place he advised boiling the drinking-water, and optionally ad- 
ding some cinnamon in polluted areas. When settled at Leiden Blume published a book on Asiatic 
cholera (A: 1831). Shortly after, he attended a congress of naturalists and surgeons at Halle, a 
town at that time suffering from a serious epidemic of cholera. He observed that in the rather 
isolated 'Franckische Stiftung', a community of some 1800 souls, there was no cholera. These 
people were followers of the pietist A.H. Francke, founder of this 'Stiftung' in 1663. To his 



satisfaction he observed that this group of people got its own water from wells through a systein 
of tubes several miles outside Halle. In Holland, where at that time cholera also was a serious 
disease, he noted that it was rare in the southwestern island province of Zeeland, and he correlated 
this with the fact that drinking-water there was mostly rainwater. The next year he wrote a pam- 
phlet (A: 1832) on the subject which he had printed in 1000 copies at his own expense. He forward- 
ed free copies to all municipalities, stressing that boiling all drinking-water was the simple remedy. 
One would expect that the arguments for this cheap advice were immediately accepted, and at least 
tested. But his opinion wascompletely overruled by the powerful voice of G.J. Mulder, a chemist 
of great influence, who declared that Blume's conclusions were nonsense and that all water from 
ditches and canals was fit for drinking and had nothing to do with the dispersion of cholera. 
Blume's role looks to me similar to the one of Semmelweiss in Vienna and his fight against 
puerperal fever. Thirty years later Blume's conclusions were of course fully accepted. 

As a civil servant Blume excelled in activity for the benefit of the country and colony, in pro- 
moting the interests of agriculture and horticulture, throughout his life. As a director of the Rijks- 
herbarium he did all he could under the circumstances, to raise it to a first-rate institution. As 
my wife (C: van Steenis-Kruseman, 1979: 37) put forward, Blume succeeded in greatly enriching 
the Rijksherbarium with important standard collections, e.g. Spanoghe (Timor), Korthals (W. 
Malesia), Forsten (Celebes), von Siebold, Textor and Burger (Japan), Sieber, Schultes, 
Cuming, Persoon, Dozy, and Molkenboer (Bryophytes). Besides this, he acquired large sets of 
duplicates from the collections of Wallich, Ecklon & Drege (Cape), and Plantae Preissianae 
(Australia). He purchased also several smaller collections from South America. 

In the preceding pages I hope to have succeeded in making it clear that the slander of which 
Blume was a victim was unfounded and can be defused by factual evidence. 

I will now proceed with some remarks on Blume's personality and his motives, as an addition 
to what already may transpire from the precedings pages. Much can be learned about this from 
his published papers. A perusal of his personal letters to his colleagues abroad will add probably 
more but this falls beyond my capacity. Another source is the opinion of third parties which can 
be found, for instance, in biographical papers. However, the latter are mostly an evaluation of 
the quantity and quality of achievements and seldom enter into personal facets. Among the 
obituaries of Blume only Goddijn (B: 1931) ventilated some well-considered remarks. 

Blume was a most intelligent person devoted to science and with a broad outlook, dedicated 
to promote the interests of his second fatherland and all its inhabitants. He pleaded for a society 
in which everyone, irrespective of race, should benefit from increasing profit. He was antagonistic 
to the idea of a 'Cultuurstelsel' ' and pleaded for a free society. 

As to his social contacts, it is difficult to ascertain much factual evidence without having access 
to his personal correspondence. His family life seems to have been happy and his wife sometimes 
shared his stays abroad. In Java he had good friends, e.g. Praetorius, Spanoghe and several 
others. As to his contacts with foreign colleagues, Blume apparently often took part in the annual 
'Versammlung Deutscher Naturforscher und Aerzte' in Germany. 

in his native country he must have had friendly relations, among them the Nees von Esenbecks 
at Regensburg. According to Roland (B: 1944) he and his wife paid in September-October 1834 
a lengthy visit to Paris where he had many friends (amongst others Decaisne, Brongniart). He 
met many prominent personalities, compared material from Java o^ Araceae, Annonaceae, etc. 
with Paris collections, bought books, acquired and bought collections and frequently stayed with 
J.E. Gay (who had very rich collections) for studying material, often together with A. Moquin- 
Tandon, the monographer of Chenupodiaceae. The latter said of Blume (B: Roland, 1944: 74): 

(I) In the Netherlands East Indies the system in which ihc local people were forced lo grow various sorts 
of crop suitable for the European market (in force mainly in Java, 1828 1890). 


Flora Malesiana 

'Je suis sorti avec M. Blume dont j'aime beaucoup la figure gracieuse, la gatte et la vitalite vrai- 
ment meridionale.' 

The fact that so many honours befell him (F) indicates that he must have enjoyed the sympathy 
of many persons abroad who took the initiative to make the proposal. In political circles in 
Holland he certainly was also appreciated; the fact that he did not succeed in building up a staff 
of collaborators for which he pleaded in vain for two decades, can be ascribed to the rather poor 
economic situation of the kingdom, unfavourable for creating permanent scientific positions. 

I believe that the later strenuous relations with his Leiden scientific contemporaries must, to 
a large part, be ascribed to feelings of envy towards his great capacities by the autocratic von 
SiEBOLD and Junghuhn, the mediocre de Vriese and the frustrated Reinwardt and Hasskarl, 
who all eagerly grasped any opportunity to damage his image. In this they were in a way assisted 
by Blume's rigid, autocratic personality. 

Unfortunately it is difficult to obtain more impartial contemporaneous information from 
neutral, disinterested parties. Among the rather neutral sources there is one, from the Swiss 
Heinrich Zollinger, who wrote an extensive diary which is now deposited in the Central Library 
at Zurich (B: Zollinger, 1841). The part of this diary relating to Zollinger's stay at Leiden, Oc- 
tober to December 1841 , was typed out and generously put at my disposal by Prof. Dr. H. Wan- 
ner, Zurich. 

Zollinger, at the suggestion of A. de Candolle, was considering a botanical-zoological ex- 
ploration of Java and wanted subscriptions from biologists, authorities, and institutes for his 
endeavour. After having obtained some in Switzerland, France, and Belgium, he came to Hol- 
land, in 1841, where Miquel gave him some hope. With his letters of recommendation he tried 
to obtain subscriptions from the Rijksherbarium and from the National Museum of Natural 
History at Leiden. Above all, he sollicited free transport for himself and his equipment to Java 
from the Dutch authorities as a contribution to his future work in the colony. In his diary Zol- 
linger gave his free opinion on several scientists he visited (Reinwardt, Temminck, Schlegel, 
DE Vriese, Ammann, Splitgerber, Schwaner, von Siebold, Korthals) (D: 8). He paid visits 
to Blume and noted about him (B: Zollinger, 1841: 25): 'Blume ist ein kleines, elegantes, 
vornehmes, lebhaftes Mannchen, das sich auf verschiedene Weise ein grosses Vermogen und eine 
grosse Reputation erworben hat. Er war sehr freundlich und zuvorkommend, gab mir Rathe aller 
Art. Ob nun im Herzen es anders aussieht, warum er so gegen mich ist, weiss ich nicht. Ich will 
das Beste denken und auf meiner Huth sein'; I.e. 29: 'Er schwatzte mir freundlich vor, wie bis 
jetzt noch kein Privatunternehmen wie meines auf Java, gelungen. Wie ich dort nichts neues mehr 
finden werde, besonders im Westen; ich miisse mich zeitig nach einer Anstellung umsehen. Aus 
dem Ganzen schien mir hervorzugehen dass er mich ganz abzuhalten oder fiir den hollandische 
Dienst zu gewinnen sucht; denn auf beide Weise kommt nichts in fremde Hande, oder im letzteren 
alles zuerst in die seinen'; I.e. 31 (summarized in English): von Siebold suggests that Blume is 
a rather tough person and reckons that Zollinger will anyway send him plants, obviously 
alluding to Blume's refusal to subscribe to a set of Zollinger's plants; I.e. 33: Blume subscribed 
to buy Lichenes from Java and offered him an iron trunk. He spent another evening in Blume's 
beautiful house, with a large library, but the trunk did not turn up. 'Blume hat fiinf hiibsche 
Kinder und eine hochgebildete Frau. Er zeigte mir seine Rumphia und andere Sachen, die auf Java 
bezug haben. Wir sprachen meist von Indien. Ich soil 3 Kisten (lebende) Pflanzen miterhalten' 
(obviously for the Botanic Gardens at Buitenzorg). At the advice of Blume he went to Mr. Ar- 
RiENS, a high official at The Hague, who suggested an audience with the Minister of the Navy, 
but Zollinger had no success; all he got was a permission to collect in the colony, antiquities ex- 
cepted. In passing, Zollinger followed Blume's advice and sollicited to be attached to the 
Botanic Gardens at Buitenzorg, but there was no vacancy at that time. Thus, Zollinger had not 
much success at Leiden, as far as botany was concerned. It remains guesswork whether Blume 
could have achieved more for him if he had backed him up. 

Summing up my impression of Blume's personality, it appears that he was not a social, 



amicable person, but self-centered and keeping aloof; also conscious of his capacities and dignity 
but lacking flexibility. However, his motives were honest, and this becomes clear from scanning 
his own writings and other literature, if judged against the background of his time and cir- 
cumstances. It is true that he had a sharp pen and in defending the rights and interests of the Rijks- 
herbarium his acid reprimanding of Junghuhn, no less a dominant authority than himself, un- 
necessarily hurt personally, which, to say the least, led to a severe estrangement. 

However, the slander to which Blume became a victim is unjustified, and may well have been 
induced by jealousy of his brilliant scientific achievements and envy of his monopoHstic position 
at the Rijksherbarium. In my view Blume was an enlightened scientist, whose image may hereby 
be restored. 

C. G.G.J. VAN Steenis 


Flora Malesiana 

Appendix A - Blume's publications 

1817 - Dissertatio inauguralis medica, de Arsenica et Ratione qua in Animalia agit. Leiden. 49 
With verses by D.J. Veegens, a friend, and Prof. S.J. Brugmans. 

1821 - Minerale wateren van Tjipannas en Tjiradjas. 
Bataviaasche Courant, 15 Sept. 1821. 

Repr. in Indisch Magazijn, Tweede Twaalftal, no 1/2, 1845: 162-166. 

Contains a chemical analysis, obviously made by Blume himself, of the mineral contents 

of these waters. 

1822 - Gedachten op eene reize door het Zuid-Oostelijk gedeeUe der Residentie Bantam. 
Bataviaasche Courant, 16 Febr. to 30 Nov. 1822. 

Repr. in Indisch Magazijn, Tweede Twaalftal, nos 3/4, 1845: 1-36. 

Report of trip, describing the history, anthropology, ethnography and poHtics of the Badui 

people in SW. Java. No botany involved. 

1 822 - Beschrijving van de heilige graven der Badoeis in het Zuid-Oostelijk gedeelte der Residen- 
tie Bantam. 

This appeared as the chapter 'Mengelingen' in the Bataviaasche Courant, nos 7, 8, 10, 13, 

27-29, and 32, 16 Febr. to 30 Nov. 1822. 

Repr. in Indisch Magazijn, Tweede Twaalftal, nos 3/4, 1845: 1-36. 

An ethnographical description of the Badui people in SW. Java, their sacred graves, etc. 

In the library of the Institute Taal-, Land- & Volkenkunde, Leiden University, there is a 85 

pp. manuscript (H 75) with the title 'Gedachten op eene reize, in de maanden December en 

Januari jl., in het zuidoostelijke gedeelte der Residentie Bantam gedaan. Getrokken uit de 

Javaansche Couranten van J 822,' which is probably copied literatim. 

1 823 - Catalogus van eenige der merkwaardigste zoo in- als uitheemsche gewassen, te vinden in 
's-Lands Plantentuin te Buitenzorg. Batavia. 112 pp., 1 pi. 

Several new genera and species. Many nomina nuda under Reinv^ardt's name. 
Repr. in Arnold Arboretum 1946. 

1823 - Beschrijving van eenige gewassen, waargenomen op eenen togt naar den Salak in den 
jaare 1822. 

Verhand. Batav. Genootschap van K. & W. 9: 129-202. 

Mostly descriptions of plants {Magnoliaceae, Loranthaceae, Dipterocarpus, Cedrela, Piper, 

1823 - Letter to Nees von Esenbeck. Flora 6: 713-716. 
Report on a planned trip in Java. 

1823 - Bijdrage tot de kennis onzer Javaansche eiken. 

Verhand. Batav. Genootschap van K. & W. 9: 203-223, 6 pi. 
Account of Quercus in Java (incl. also Lithocarpus). 

1823 - (with C.G. Nees von Esenbeck) Pugillus plantarum Javanicarum, e Cryptogamicarum 

variis ordinibus selectus. 

Nova Acta Acad. Caes. Leop. -Carol. 11 (1): 117-138, pi. 12 & 13. 

Descriptions of Pteridophytes, the new species under dual authorship 'Nees & Bl.'. 


1 824 - Letter to Nees von Esenbeck: Veber die Vegetation des Berges Gedee aufder Insel Java. 
Flora 7: 289-295. 

Extract from a larger paper in Dutch, see below (1825). Sketches on the exploration of Mt 
Gedeh made together with the hortulanus Kent. Blume did not ascend Mt Pangrango. 

1824 - Epidemie onder de buffets. 

Bata\iaasche Courant, 10 Jan. 1824: 'Verslag van den kommissaris van den burgerlijk 

geneeskundigen Dienst in Nederlandsch Indie C.L. Blume.' 

See also: Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 91-94. 

Epidemic disease among the buffaloes. 

1825 - Letter to the Governor-General, dated 8 Dec. 1824, published in the Bataviaasche 
Courant, 12 Jan. 1825. 

Report on Blume's discovery of Rafflesia in Nusa Kambangan I. (S. Java), the first 
discovery of the genus in Java. He did not name it here. 

1825 - Bestijging van den berg Tjerimai, gewoonlijk genoemd Tjerme, in de Residentie Cheri- 

Bataviaasche Courant, 2 Febr. 1825. 

Repr. in Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 102-116. 
Report of a trip from Krawang eastwards to Panarukan, Linggadjati, culminating in the as- 
cent of Mt Tjeremai, with many botanical data on plants encountered. 

1825 - Over de gesteldheid van het gebergte Gedeh. 

Verhand. Batav. Genootschap van K. & W. 10: 55-104. 

Lively topographical and botanical description of an ascent of Mt Gedeh from Bogor via 
Puntjak, along Megamendung, Tjibeureum and Kandangbadak through the crater and 
along the Alun-Alun to the summit. Blume did not ascend Mt Pangrango, and thus missed 
Primula imperial is. 

1 825 - Inlandsche middelen tegen diarrheen. 
Bataviaasche Courant, 23 Febr. 1825. 
Native recipes against diarrhoea. 
See also: Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 116. 

1825 - Tabellen en Platen voor de Javaansche Orchideen. Batavia. 5 tab., 16 pi. Folio. 

Famous exposition of a system of the Javanese orchids and their affinities; 73 spp. depicted 
in detail. Issued with the Bijdragen (1825-1827) part 6. 

1825 - (with C.G. Nees von Esenbeck & C.G.C. Reinwardt) Hepaticae Javanicae editae con- 
junctis studiis et opera. 

Nova Acta Acad. Caes. Leop. -Carol. 12: 181-238, 409-417. 
Account of hepatics in Java. 

1825 - lets over de planten onder den naam van Paima. bij de Hindostaners en de Javanen 

Bataviaasche Courant, 9 March 1825. 

Repr. in Indisch Maga/ijn, Tweede Twaalftal, no 3/4, 1845: 179-183. 
A note on plants known under the vernacular name 'patma' ( = Rafflesia). 


Flora Malesiana 

1825 - Korte beschrijving van de Pat ma der Javanen. 

Bataviaasche Courant, 23 March 1825 (22 pp., in L). 

Repr. in Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 183-194. 

Short description of the 'patma' (-Rafflesia) of the Javanese. 

1825 - Die Patma-Pflanze der Indier und Javanesen und Beschreibung einer neu entdeckten 
Blume auf der Insel Noesa Kambangan, die an Grosse alle bis dahin bekannt gewesenen 

Liter. Wochenbl. der Borsenhalle, Hamburg, no 29: 454-462. Repr. in L. 
As the preceding. 

1825 - Beitrage zur Kenntnis von Bantam, dem westlichsten Bezirk auf Java. 
Hertha II: 227-257. 
Not seen. Probably similar to entries in 1822. 

1825 - Letter to Th.F.L. Nees von Esenbeck: Reise von Batavia nach Krawang in der Preanger 
Regentschaft. Flora 8 (2): 577-585. 
Report of journey from Batavia to Krawang. 

1 825 - Etwas Uber die Rhizantheae, eine neue Pflanzenfamilie, und die Gattung Rafflesia insbe- 
Flora 8 (2): 609-624. 

1825 - Letter to Th.F.L. Nees von Esenbeck: Ueber Pflanzen der Gegend von Batavia. Flora 
8 (2): 676-680. 
Flora of the vicinity of Batavia. 

1825 - Letter to the Governor-General, dated 20 Nov., on the flowering of a new species of a 
new genus of Araceae with a very large inflorescence, obviously Amorphophallus cam- 
panulatus, in the Botanic Garden, with reference to Tacca phallifera Rumph. 
Bataviaasche Courant, 23 Nov. 1825. 

1825-1827 - Bijdragen tot de Flora van Nederlandsch Indie. 17 fascicles, 1169 pp. 

For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 236. 

In all 107 families are treated, in which 700 genera and over 2300 species were incorporated. 

There are many new genera and very many new species, all described in concise Latin. In 

the first 5 fascicles each family has also a paragraph with notes on its useful plants. 

On p. 265 Blume mentioned that his plan was to treat the orchids together with van 

Hasselt; 27 species out of the 296 were jointly described. Through van Hasselt's early 

death this joint venture was frustrated. 

There is a typed Index to the names in L. 

Data on useful plants mentioned in fascicle 1 were copied in Alg. Konst- en Letterbode 1826- 

1: 26-29, 37-41. 

1826 - Monographie der Oost-Indische pepersoorten. 

Verhand. Batav. Genootschap van K. & W. 11: 139-245, 6 pi., 41 fig. 
Monography of Netherlands-Indian species of Piper. 

1826 - De Tacca Culat van Rumphius wedergevonden. Mededeeling van de waarnemingen van 
C.L. Blume. 
Alg. Konst- en Letterbode 1826-1: 333-334. 



Report about Blume's recollection of a Rumphian aroid in the island of Nusa Kambangan, 
S. Ja\a: Amorphophallus campanulatus. 

1826 - Letter to Nees von Esenbeck: Bruchstucke einer Reise auf der Insel Java. Flora 9 (2): 
417-426, 433-441. 

Report on a trip in NW. Java, including also an ascent of Mt Tjeremai. 

1827 - (with Th.F.L. Nees von Esenbeck) Fungi Javanici. 
Nova Acta Acad. Caes. Leop. -Carol. 13 (1): 9-22, pi. 2-7. 

1827 - Over een nieuw plantengeslacht, de Brugmansia, uit de natuurlijke familie der Rhizan- 

In: H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen 2: 419-423. 
Brugmansia, a new genus of the Rafflesiaceae. 

1 827 - Observations sur le structure des poivres. 
Ann. Sc. Nat. 12: 216-224. 
Extract in French of the monograph of Piper (1826). 

1827 - Bijdrage tot de kennis van het landschap Bantam, in het westelijk gedeelte van Java, etc. 
Cybele (Tijdschr. Bevordering Land- en Volkenkunde) VI' stuk: 1-36. 
Contribution to the knowledge of Bantam, West Java. Almost literatim reproduced under 
the same title in Indisch Magazijn, Tweede Twaalftal, no 3/4, 1845: 1-36. 

1 827 - Over de siaat der indigo-teelt. 

In: P. VAN Griethuizen, Over de staat der indigo-teelt. De Nederl. Hermes, Tijdschr. 
Koophandel, Zeevaart en Nijverheid 2, no 10: 40-42. 
Brief information and references on cultivation of indigo. 

1827-1828 - Enumeratio plantarum Javae et insularum adjacentium minus cognitarum vel 
novarum ex herbariis Reinwardtii, Hasseltii, Kuhlii, Blumei, etc. Leiden. 2 vols. 278 pp. 
Description of some families of Angiosperms and the Pteridophytes. Properly a continua- 
tion of the Bijdragen (1825-1827), although in more detail and with longer descriptions. 
Repr. Den Haag 1830, Amsterdam 1968. 

1 828 - Het Duizend-Gebergte (Goenong SeribuJ. 

In: G.H. Nagel, Schetsen uit mijne Javaansche portefeuille; Javaansche tafereelen: 69-75 
(in L). 

Remarks on the landscape of the 'Thousand Hills', in the plain SW of Jakarta. Also a brief 
description of the limestone hills Kuripan, SW of Bogor, famous for their hotsprings, which 
yielded several plants not found anywhere else, amongst them a Cycas sp. 

1828-1851 - Flora Javae nee non insularum adjacentium. Brussels. 3 vols. 

Three sumptuous folio volumes, with analyses, plates, and descriptions in great detail. The 
authorship is partly ascribed to his assistant Dr. J.B. Fischer, who was his 'adjutore'. The 
preface is probably most interesting, but being not in sufficient command of the Latin 
language, 1 cannot evaluate it. 

For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, I (1976) 236. 
23 Planches in^dites were for sale in probably 1863 (see also C: van Steenis, 1947). 

1829 Letter to the Governor-Cicncral. Algcmccn Handelsblad of April Isi, no 26. 


Flora Malesiana 

On the occasion of the appointment of Governor-General van den Bosch; on the impor- 
tance of stimulating cultures for the general welfare, commerce, and the benefit of the com- 
mon people. Blume pleaded for the gradual abandoning cq. restriction of the use of opium. 

1831 - Reistogte naar Buitenzorg, het Duizend-Gebergte, Koeripan en in de omstreken van 
Batavia, 1824; door een ambtenaar. 

Recensent (de Recensenten) XXIV, 2: 427-442, ^dl-AlX. 

This contribution is not written by Blume himself, but by one of the civil servants accom- 
panying him, A. ZipPELius or A. Latour, on a trip to the hills W of Bogor. By Blume 
himself also described in the entry of 1828, Het Duizend-Gebergte. 
Contains no scientific observations. 

1831 - Ueber einige Ostindische, und besonders Javanische Melastomataceen. 
Flora 15 (2): 465-527. 

A thorough study of the family Melastomataceae in which Blume described 12 new genera, 
all standing to the present day, mainly based on species described in the Bijdragen 

1831 - Over eenige Oost-Indische, byzonder Javaansche, Melastomataceae. 

In: H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen 6: 211-268. 
The same as the preceding entry. 

1831 - Eenige woorden over de redding van het Rijks Herbarium door Dr. J.B. Fischer. 
Alg. Konst- en Letterbode no 23, 10 June: 356-359 & no 24, 17 June: 314-311 (in L). 
Detailson the transfer of the Rijksherbarium from Brussels to Holland by Dr. J.B. Fischer. 

1831 - Over de Asiatische cholera, uit eigene waarnemingen en echte stukken. C.G. Sulpke, 
Amsterdam, viii + 203 pp. (In University Library at U). 

Historical account and personal experience with cholera in the Netherlands Indies, exten- 
sively documented; measures taken by the government to cope with this disastrous illness. 

1 832 - Vruchten mijner ondervinding in het afweren en genezen der cholera. Amsterdam. 3 1 pp. 
(in L). 

A most interesting paper prescribing how to deal with patients suffering from cholera, in 
Java caWedfebris endemica bataviae. Recipes for external and internal use. Prescribing the 
boiling of drinking-water. Paper printed in 1000 copies at the author's expense, distributed 
freely to boards of municipalities in the Netherlands. 

1832 - Beschrijving van Calamus draco Willd., etc. 

In: H.C. VAN Hall (ed.), Bijdragen tot de Natuurkundige Wetenschappen 7: 115-129. 
Extensive Latin description of a rattan from S. Sumatra collected by his friend C.F.E. 

1832 - Uittreksel uit eenen brief van den Heer J.B. Spanoghe aan den hoogleeraar C.L. Blume. 
Alg. Konst- en Letterbode 1832-1: 356-361. 

Notes on the situation in Bima (Sumbawa), with biographical notes on Spanoghe by Blume. 
Plant list of Bima. 

1834 - Observationes de genere Helicia Lour. 
Ann. Sc. Nat. ser. II, 1 (1) Bot.: 211-220. 
Review of the genus, with new species. 



1834 - Eenige waarnemingen omtrent de Culilawan boom van Rumphius. 
Tijdschr. Natuurlijke Geschiedenis en Physiologie 1: 45-64, t. 2. 

Repr. in: Wiegman, Archiv Naturgeschichte 1 (1835) 1 16-126, and in: Jahrb. Pharm. Berlin 
35 (1835)9-29. 
On Cinnamomum described by Rumphius. 

1834 - Eenige opmerkingen over de natuurlijke rangschikking van Rohdea, Tupistra en 
Aspidistra, als mede de beschrijving eener nieuwe soort van dit laatste geslacht. 
Tijdschr. Natuurlijke Geschiedenis en Physiologie 1: 67-85, pi. 3 & 4. 
Botanical relations between three genera, and description of a new species of Aspidistra. 

1834 - De novis quibusdam plantarum familiis expositio et olim jam expositarum enumeratio. 
Tijdschr. Natuurlijke Geschiedenis en Physiologie 1: 131-162. 

Repr. in Ann. Sc. Nat. sr. II, 2 Bot.: 89-106. 

A preprint was issued in 1833, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 


Description of a number of newly proposed families, Apostasiaceae, etc., with a few new 


1835 - Neesia, genus plantarum javanicum repertum, descriptum et figura illustratum. 
Nova Acta Acad. Caes. Leop. -Carol. 17 (1): 73-84, pi. 6. 

A new genus of Bombacaceae named after Th.Fr.L. Nees von Esenbeck. 

1835-1848 - Rumphia, sive commentationes botanicae imprimis de plantis Indiae orientalis, 
tum penitus incognitis turn quae in libris Rheedii, Rumphii, Roxburghii, Wallichii, aliorum, 
recensentur. Leiden, Amsterdam. 4 vols. Folio. 

For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 238. 
Conditions for sale were mentioned in Ann. Sc. Nat. ser. II, 4 (1835) 318. 
JuNGHUHN mentions that part of the work was elaborated and illustrated by J. Decaisne 
and the Latin was supervised by D.J. Veegens, a friend of Blume. 

1837 - Levensbyzonderheden van Franz Junghuhn. 
Alg. Konst- en Letterbode 1837-11: 277, footnote. 
Biographical notes on F. Junghuhn. 

1837 - Levensbyzonderheden van Dr A. Fritze. 

Alg. Konst- en Letterbode 1837-11: 277, footnote. 

Biographical notes on A. Fritze, Inspector of Physicians and benefactor of Junghuhn. 

1837 - Naschrift op den brief van Junghuhn uit Djocjakarta. 
Alg. Konst- en Letterbode 1837-11: 278-280. 
Appendix to a letter of Junghuhn. 

1838 - Revue des palmiers de I'archipel des Indes orientates. 
Bull. Sc. Phys. & Natur. en Neerlande no 9: 61-67. 
Repr. in Ann. Sc. Nat. ser. 11, 10 Bot.: 369-377. 

1838 - Miquetia, genus novum plantarum javanicarum. 
Bull. Sc. Phys. & Natur. en Neerlande no 13: 93-95. 
Repr. in Ann. Sc. Nat. sdr. II. 10 Bot.: 255-256. 


Flora Malesiana 

Description of a new genus of Icacinaceae, named after Miquel, then director of the Rotter- 
dam Botanical Garden. 

1838 - (transl.) Advertisement for sustaining the edition of Flora Javae, Rumphia, etc. 
Alg. Konst- en Letterbode 1838-11: 322, 401. 

1839 - Beschrijving der minerale bronnen, welke nabij Tjiratjas in de Residence Krawang 
warden gevonden. 

Tijdschr. Ned. -Indie 2 (1): 451-455. 

Description of mineral wells near Tjiratjas in Krawang, E of Jakarta. 

1843 - Levensbyzonderheden over Th.St. Raffles. 
De Indische Bij 1: 49, footnote. 
Praise of Raffles' humane government. 

1843 - Engeland's staatkunde omtrent China. 
De Indische Bij 1: 61-77. 

To stimulate the necessity of increasing naval power in Netherlands-Indian waters and ex- 
tend commercial relations with Japan. In a footnote on p. 76 Blume refers again to the 
necessity of regulating the trade in opium. 

1843 - Toelichting aangaande de nasporingen op Borneo van G. Miiller. 
De Indische Bij 1: 103-176. 

On the geography, anthropology, commercial situation etc. of W. Borneo, from cor- 
respondence with G. MiJLLER. In a footnote on p. 104 Blume reveals the bad management 
of the Governor-General Daendels, and he praises Raffles for his humane, unselfish ad- 

1843 - Bladvulling. 

De Indische Bij 1: 320. 

An occasional note on common social progress, whereby also the native people should pros- 
per. Private property of land by non-natives is discouraged. Native rule should not be under- 
mined. Adat should be maintained. 

1843 - Over een Nederlandsch Gezantschap in Japan. 
De Indische Bij 1: 479-480. 
Importance of a Netherlands Embassy in Japan. 

1843 - Over eenige Oost-Indische planten welke eene uitmuntende vezelstof opleveren, en 
Gedachten over het nut van dergelijke kulturen tot opbeuring van de buiten Java gelegene 

De Indische Bij 1: 481-509. 

On the importance of fibres, from ramie, cotton and Musa; tissues provided by Blume were 


1844 - (with P.P. VON Siebold) Ontwerp tot oprigting van de Koninklijke Nederlandsche 
Maatschappij tot Aanmoediging van den Tuinbouw. 

Jaarb. Ned. Mij. Aanmoed. Tuinbouw over 1844: iii-iv. 
Tentative rules for the newly erected society. 



1844 - Over het nut der invoering van vreemde gewassen en de laatste pogingen om daardoor 
den tuinbouw hier te lande op le beuren. 
Jaarb. Ned. Mij. Aanmoed. Tuinbouw over 1844: 41-88. 
On the use of importing exotic plants for horticulture in the Netherlands. 

1 844 - Naamlijst van Oost-Indische en bepaaldelijk Javaansche gewassen, etc. 
Jaarb. Ned. Mij. Aanmoed. Tuinbouw over 1844: 88-90, t. 1-4 (col.). 
Unsigned, but attributed to Blume. 

1 844 - Ueber das Lycopodium arboreum Jungh. 

Amtlicher Bericht uber die Versammlung Deutscher Naturforscher and Aerzte Abt. 2, 22: 


Identified as Dacrydium cf. elatum Wall, on type material shown to him by W.H. de 

Vriese. In Rumphia 3 (1849) 219, 221 Blume later added sour remarks. 

1 844 - Ueber ein Surrogat des Chinesischen Thees. 

Amtlicher Bericht iiber die Versammlung Deutscher Naturforscher und Aerzte Abt. 2, 22: 


Made public in a session of the Society at Bremen, 23 Sept. 1844. As Prof. G.J. Mulder 

had shown the alkaloid theine is the same as caffeine, Blume suggested that tea could be 

made from dried leaves of coffee. 

1845 - De Koffij-thee. 

Astrea, Tydschr. van Schoone Kunsten, Wetenschap en Letteren 1: 285. 
Same as preceding. 

1 845 - Minerale wateren van TJipannas en Tjiradjas. Opmerkingen nopens de bruikbaarheid van 
dien le TJipannas (Preanger Reg.), beschrijving en scheikundig onderzoek van dien te 
Tjiradjas (Krawang). 

Indisch Magazijn, Tweede Twaalftal, no 1/2: 162-166. 
Reprint of an article published in the Bataviaasche Courant of 15 Sept. 1821. 

1845 - Gedachten op eene reis door het zuidoostelijk gedeelte der Residentie Bantam. 
Indisch Magazijn, Tweede Twaalftal, no 3/4: 1-36. 

Account of his experience on a trip through SE. Bantam in W. Java. Account of the Badui 
people. Reprint of an article published in 1822. 

1845 - Fragment uit een Dagboek gehouden op eene reis over Java. Bestijging van den berg 
TJerimai, gewoonlijk genoemd Tjerme, in de Residentie Cheribon. 
Indisch Magazijn, Tweede Twaalftal, no 3/4: 102-116. 

Report on an exploration of Ml Tjeremai, above Cheribon. Reprint of an article published 
in 1825. 

1845 - Over iniandsche middelen tegen diarrhoe. 

Indisch Magazijn, Tweede Twaalftal, no 3/4: 1 16. 

Indigenous recipes against diarrhoea. Copied from the paper published in 1825. 

1845 - De patma van Noesa Kambangan. 

Indisch Magazijn, Tweede Twaalftal. no 3/4: 179-194 (in L)- 

Reprint of an article published in the Bataviaasche Courant of 9 & 23 March 1825, in which 
he described his finding of Rafflesia in Nusa Kambangan I. (S. Central Java) and claimed 
this to be the largest flower, superseding Nelumbntm. 


Flora Malesiana 

1846 - An article in the 'Handelsblad'. 

In this article Blume advised to hold expositions of colonial products from the East and 
West Indies in the Netherlands from time to time. 

1849-1856- Museum botanicum Lugduno-Batavum sive stirpium exoticarum, novarum vel 
minus cognitarum ex vivis out siccis brevis expositio et descriptio. Leiden. 2 vols. 
Appeared in dated parts each of 16 pp. In all, 1: 396 pp., 60 fig.; II: 256 pp., 58 fig. The 
second volume was not finished and had no index. 

For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 240. 
A most important work, being a scientific catalogue of the Rijksherbarium collections, 
hence containing descriptions and treatments of plants from all over the world. 
An Index to volume 2 was prepared by C. G.G.J, van Steenis & Chew Wee Lek at the 
Rijksherbarium in 1974 (see Appendix C). 

1850 - Antwoord aan den Heer W.H. De Vriese. 

Alg. Konst- en Letterbode 1850-11: 99-109, 114-123. Repr. 34 pp. in L. 
Blume defends his criticism on the identity of Lycopodium arboreum and the reduction of 
Pinusmerkusii, and the right of the Rijksherbarium to be the depository of collections made 
by civil servants. 

1850 - Opheldering van de inlichtingen van den Heer Fr. Junghuhn. 

Alg. Konst- en Letterbode 1850-11: 258-261, 274-279. Repr. 19 pp. in L. 
On Junghuhn's collection and the right of the Rijksherbarium as the proper public deposi- 
tory of botanical collections. 

1852 - Copy of a letter to J.G. Baud, Minister of the Colonies, dated 14 March 1840, 'nopens 
de bereiding van thee uit koffie-bladeren, met aanbeveling tot het nemen van proeven in het 
groot op Java zelf. ' 

Natuurk. Tijdschr. Ned. Indie 3: 122-126. 

Proposal to prepare tea from coffee leaves and suggesting experiments with this on a large 
scale in Java. 
There are two other entries on the subject in 1844 & 1845; see also Astrea 1 (1851) 256. 

1855 - Melanges botaniques. 8°. No 1, 1 Aug. 1855: 1-8; no 2, 1 Sept. 1855: 9-12. Facsimile 
in Taxon 35 (1986) 274-285. 

Until June 1985 assumed not to have been published; see Stafleu & Cowan, Taxonomic 
literature, ed. 2, 1 (1976) 241. 

The new names etc. in the Melanges were validated by Walpers in his Annales 4 (1857) 
6A1-6AA and a rather large extract was published in Flora 41 (1858) 254-256. 
L. Vogelenzang, librarian of the Rijksherbarium, found in Vesque's bibliography of 
J. Decaisne (C: 1883) that the latter had a copy of the Melanges in his library, now incor- 
porated in the Bibliotheque Nationale at Paris. H. Heine located another copy in the 
Bibliotheque Central of the Museum d'Histoire Naturelle at Paris which had belonged to 
the library of A.Th. Brongniart. The original copy mentioned in Flora is still not located. 
It was probably dedicated to Nees von Esenbeck. 

The pamphlet was not for sale, but it was effectively published and at least two copies exist. 
Both Paris copies were autographed to Blume's close friends. He may have sent more copies 
to other botanists with whom he was befriended. Obviously Blume published it at his own 
expense and the reason for this is unknown. He could have published it in his Museum 
Botanicum Lugdunum-Batavum . 
The first numero of the Melanges contains a discussion on paper-making by the Sino- 



Japanese and three species are described of Broussonetia (2 new). Furthermore there is a sec- 
tion 'synonymic de quelques plantes peu connues', concerning species and genera of Cuno- 
niaceae, Saxifragaceae, Rosaceae, Guttiferae (Cratoxylon), Dipterocarpaceae, Ulmaceae, 
Moraceae, and Nepenthes. Numero 2 contains Chrysobalanaceae and Rosaceae (Pygeum) 
(B: VAN Steenis, 1986). 

1858 - Bijdrage tot de kennis der Oost-Indische Orchideen en het maaksel (de organisatie) van 
hare bevruchiingswerktuigen. 

Versl. & Meded. Kon. Ned. Akad. Wetensch., Amsterdam 7: 100-115, 2 pi. 
Interpretation of the orchidaceous flower, with special regard to Apostasiaceae. 

1858(-1859) - Flora Javae et insularum adjacentium. Nova Series. Leiden, pp. 8 + 6+ 162, 66 
col. pi. 

Also edited with a French title, see below. 

A sumptuous work in which Blume summarized his large knowledge on orchids in which 
he had great insight since he wrote the Bijdragen (1825-1827). 

1 858(- 1 859) - Collection des Orchidees les plus remarquables de I'Archipel Indien et du Japon. 
The French-titled version of the Flora Javae, Nova Series. 
For publication dates, see Stafleu & Cowan, Taxonomic literature, ed. 2, 1 (1976) 240. 

1859 - (with A.H. van der Boon Mesch) Geschikte materialen uit de Overzeesche bezittingen 
voor het vervaardigen van papier. 

Report about useful materials from overseas territories suitable to manufacture paper. 

1859 - Vanda suaveolens Bl. 

Ann. Hort. Bot. ou Fl. Jard. Pays-Bas 2: 1-2, 1 col. pi. 

1859 - Over eenige Oost-Indische houtsoorten in verband met de verwoestingen door den paal- 
worm of andere schelpdieren hier te lande en elders aangerigt. 

Versl. & Meded. Kon. Ned. Akad. Wetensch. 9: 25-49. Repr. 25 pp. in L. 
A scholarly review of timbers resistant against teredo and other molluscs, in which Blume 
summarized experience onwards of Rumphius and collected data from all kinds of sources, 
indicating valuable species to be used in sea harbours. 

1860 - De houtteelt verbonden met den landbouw. 

Tijdschr. Ned. Mij. ter Bevordering van de Nijverheid 23: 1-29. 
Cultivation of timber species in relation to agriculture. 

1861 - Monographie des Anoectochilus, Goodyera et genres voisins, les plus remarquables de 
I'archipel Indien et du Japon. 

Belg. Hon. 11: 369-378, 1 pi. 

Extract from Flora Javae, Nova Series (1858-1859). 

1863 - Flora Javae. Planches in^dites. 

23 coloured folio plates of Javanese plants with names and analyses. These were probably 

intended for further instalments of the Flora Javae, but remained without text. 

On the back of some plates an advertisement was printed by a booksellers firm in Leiden; 

herein Blumf.'s works were offered for sale, as a packet, probably one or (wo years after 

his death. 

I have distributed a few copies to some herbaria, with a nolo, in November 1947 (C: 1947). 

Further particulars I published in Blumea 6 (1948) 263. 


Flora Malesiana 


1 823 (April) - Herinnering aan acht merkwaardige dagen van mijn /even, op een uitstapje naar de 
top van de Gounong (berg) Gedu. 

This concerns a 16 pp. manuscript which has wrongly been attributed to Blume. It was writ- 
ten by a party following Blume's trail to the lower part of the crater of Mt Gedeh above 
Tjibodas. It is preserved in the library of the Instituut van Taal-, Land- en Volkenkunde, 
Leiden University (H 338). 

Appendix B — Biographical sources 

Aa, A.J. VAN DER. 1878. Biographisch woordenboek der Nederlanden. Bijvoegsel: 34-35; ibid.: 

111-115. - A concise biography. 
Anonymous. 1827-1856. Algemeene Konst- en Letterbode 1827-11: 137; 1829-1: 227; 1831-1: 50, 

359; I833-I: 429; 1838-11: 290; 1851-1: 257; 1853-1: 193, 305; 1855: 118; 1856: 57. 
Anonymous. 1853. Bonplandia 1: 228. - Blume was in Berlin and offered (obviously at a meet- 
ing) fibres of Boehmeria tenacissima Bl. which he said had a great durability and could possibly 

be of importance for the navy. He was then presented to the King of Prussia. On the fibres of 

this Boehmeria he pubUshed in the Melanges botaniques (A, 1855). 
Anonymous. 1855. Bonplandia 3: 155. - Here it was reported that Reinwardt sold his library 

for Dfl. 20,000. His herbarium was donated to the University herbarium of Leiden, on the con- 
dition that it should not be incorporated in the Rijksherbarium. 

N.B. In the 'Instruction' of 1832 (see C: van Dam) it had been officially decreed that the 

University herbarium was to be merged with the Rijksherbarium! 
Anonymous. 1858. Flora 41: 254-256. - Extract review oi Melanges botaniques. 
Anonymous. 1862. Leidsch Dagblad, 5 Febr. 1862, no 598. Repr. of 3 pp. in L. - Formal 

Anonymous. 1862. Bonplandia 10: 47. - Obituary note. 
Anonymous. 1862. Botanische Zeitung 20: 56. - Obituary note. 

Anonymous. 1862. Proceedings Linnean Society of London 1862: xcvi-xcviii. - Obituary note. 
Anonymous. 1862 or 1863. Annuaire de 1' Academic de Paris. - Obituary note (not seen). 
Anonymous. 1875. Allgemeine Deutsche Biographic 2: 746-747. - Short biography. 
Anonymous. 1875. Album Studiosorum Lugdunum Batavum 1575-1875, column 1243. - Short 

Anonymous. 1930. Nieuw Nederlandsch Biographisch Woordenboek 8: 132-133. - Short 

Backer, C.A. 1936. Verklarend woordenboek van wetenschappelijke plantennamen: 70. - Brief 

Baillon, H.E. 1877. Dictionnaire de Botanique 1: 433. 
Beumee, J.G.B. 1948. C.L. Blume, Museum Botanicum. Fl. Males. Bull, no 3: 69-70. - On the 

dates of publication. 
Boerlage, J.G. 1896. Botanische literatuur. Encyclopaedic van Nederlandsch-Indie ed. 1, 1: 210, 

272-273, 280. 
Bretschneider, E. 1898. History of European botanical discoveries in China: 308-309. London. 

- Brief biography; Blume illustrated some Chinese plants. 
BuRDET, H.M. 1972. Cartulae ad botanicorum graphicem. Candollea 27: 327-328. 
Candolle, a. DE.1862. Memoires et souvenirs de A. P. de Candolle: 150, 383, 412. 
1880. Phytographie: 318. - Praises the excellent figures in Blume's Museum Botanicum. 



CoLENBRANDER, H.T. 1926. Kolonialc Geschiedenis 3: 111. 

Danser. B.H. 1938. Who can give further information about the dates of publication of Blume's 
Flora Javae? Chron. Bot. 4: 454-455. 

1939. The publication dates of Blume's Flora Javae. Blumea 3: 203-211. 

GoDDiJN, W.A. 1931. 's-Rijks Herbarium 1830-1930. Meded. Rijksherb. 62b: 1-53. - Rather 

extensive biographical notes. 
GiJZEN, A. 1938. 's-Rijks Museum van Natuurlijke Historie, 1820-1915: 100-101. Rotterdam. 

- On Blume's zoological contributions to the Leiden Museum. 

Hall, H.C. van. 1862. C.L. Blume. De Nederl. Spectator, 22 Febr. 1862, no 8: 57-59. - 
Biographical data; rather extensive (in L). 

Hasskarl, J.K. 1850. Antwoord aan den heer C.L. Blume, wegens onderscheidene te mijnen aan- 
zien geuite beschuldigingen, vervat in zijn antwoord aan den heer W.H. de Vriese, Leiden 1850. 
Alg. Konst- en Letterbode 1850. Repr. of 16 pp. in L. - Hasskarl defending his rights to have 
a private herbarium. 

Jacobs, M. 1980. C.L. Blume (1796-1862). Fl. Males. Bull, no 33: 3362-3363. 

Jansen, p. & W.H. Wachter. 1941. Ned. Kruidk. Arch. 51: 343. - Biographical references. 

JuNGHUHN, F. 1837. Brief aan C.L. Blume vanuit Djocjakarta, 2 Febr. 1837. Alg. Konst- en Let- 
terbode 1837-11: 275-277. 

1850. Inlichtingen aangeboden aan het publiek over zeker geschrift van den heer C.L. 

Blume, en antwoord aan dien Heer. Alg. Konst- en Letterbode 1850, no 41. Repr. 9 pp. in L. 

- Self-defense in keeping his private herbarium. 

1850. Vervolg der inlichtingen aangeboden aan het publiek over een geschrift van den heer 

C.L. Blume. Alg. Konst- en Letterbode 1850. Repr. 29 pp. in L. - Polemics with Blume. 

1851. Een woord over den Sambinoer-boom van Sumatra, betrekkelijk deszelfs botanische 

bepaling. Ned. Kruidk. Arch. 2: 2-16. - On Blume's reduction of Junghuhn's Lycopodium 
arboreum to Dacrydium. 

1853. Java, zijnegedaante, zijn plantentooi, en inwendigebouw, 1: 183-186. 2nd Dutch ed. 

Kalkman, C. 1979. The Rijksherbarium, past and present. Blumea 25: 13-26, especially p. 14. 
KosTER, J.Th. Facsimile handwritings of Blume. - Unpublished (in L). 

Lasegue, A. 1845. Musee botaniquede M. Benjamin Delessert: 268,293,307,315,346,347,506, 

535, 562. 
Leenhouts, P.W. 1980. Het Botanisch Kabinet te Franeker: 34. 
Lintum, C. te. 1913. Een eeuw van vooruitgang, 1813-1913. Zwolle (not seen). - Blume was 

far ahead of his time in having found the solution of the combat against cholera by the simple 

boiling of drinking-water. 
Maclean, J. 1979. Carl Ludwig Blume and the Netherlands East Indies. Janus 66: 15-29. - 

Period 1820-1831; valuable biographical essay. Maclean traced many letters in the Colonial 

Archives of the 'Rijksarchief, The Hague. 
Miouel, F.A.W. 1856. Review of Blume, Museum Botanicum. Bot. Zeit. 14: 185-188, 540-541. 

- MiQUEL complained severely about Blume's antedating issues of the Museum Botanicum and 
his attempts to withhold information from his colleagues. 

Guden, a. den. 1979. C.L. Blume, pcriodc 1826-1832. Unpublished essay, made under supervi- 
sion of Dr. P. Smit, Biohistorical Institute, Utrecht. - A thorough account, largely based on 
official letters and documents of the period mentioned, as present in the 'Rijksarchief, The 

Pritzel. G.A. 1872. Thesaurus litcraturac boianicae: 29. - Blumi's selected bibliography. 

Pi;lle, a. a. 1917. Botanische literaluur. lincyciopacdic van Nedcrlandsch Indie cd. 2, vol. I: 
317, 394-395; ibid. 1919. Vol. 4: 422. 

Raalten, G. van. 1825. Unpublished letter to J.G.S. van Breda (?). - Erroneous accusation that 
Blume stole property or information from Kuml & Van Hasshlt (in L). 

koiANi>. M. 1944. Alfred Moquin-Tandon. Un naturalistc ^ Paris sous Louis-Philippe. Journal 


Flora Malesiana 

d'un voyage inedit (1834). Paris, Mercure de France ed. 3: 351 pp. — Historically a most in- 
teresting booklet full of biographical data of French botanists. Blume paid a prolonged stay 
to Paris in Sept. /Oct. 1834. 
RoMER, L.S.A.M. VON. 1921. Historische Schetsen. Batavia. 335 pp., 109 pi.; a very brief 
obituary on p. 193. - It is most peculiar that in the brief history of cholera (pp. 232-238) the 
author, himself a physician, makes no mention at all of Blume's important work on the sub- 

ScHOUTE, D. 1937. Occidental therapeutics in the Netherlands East Indies during three centuries 
of Netherlands settlement. Publication of the Netherlands Indies Health Service: 114-119. - 
Cited the governmental regulations and instructions for the native chiefs, extension of the vac- 
cination, etc. Some of these might have actually been written by Blume, who was chief of vac- 
cination and later even chief of the medical service. 

Seemann, B. 1863. Journ. Bot. 1: 64. - Short obituary. 

Seemann, W.E.G. 1859. Bonplandia 7: 52-53. - Blume complained that the Netherlands 
Government did not contribute funds towards the publication of the Flora Javae, Nova Series, 
and that this was printed at his own expense. Seemann had received the volume, or at least first 
sheets of it, on 3 Nov. 1858. He criticizes Blume for having given too little attention to the works 
of LiNDLEY and Reichenbach. 

SiRKS, M.J. 1915. Indisch Natuuronderzoek: 109-112, portr. Amsterdam. - Brief biographical 

Smit, p. 1979. The Rijksherbarium and the scientific and social conditions which influenced its 
foundation. Blumea 25: 5-11. - In this excellent essay on the foundation of the Rijksher- 
barium Smit erroneously mentioned (p. 9) that Blume transferred the Kuhl & Van Hasselt 
specimens to Leiden in 1826. 

& R.J.Ch.V. ter Laage (eds.). 1970. Essays in biohistory. Regnum Vegetabile 71. 

Stafleu, F.A. 1966. Wentia 16: 28-31. - In an excellent biography of Miquel some notes on 

1970. The Miquel-Schlechtendal correspondence. A picture of European botany, 1836- 

1866. In: P. Smit & R.J.Ch.V. ter Laage, Essays in biohistory. Regnum Vegetabile 71: 
295-341. - Many data on Blume and his works. Page 307: Decaisne made several drawings 
for Rumphia. Page 324: Junghuhn sold his herbarium to the University of Leiden on the condi- 
tion that it should not be incorporated in the Rijksherbarium. Page 326: Reference to Miquel, 
who was glad that in February 1851 a new, more 'liberal' Instruction for the Rijksherbarium 
was issued by the Government. Page 33 1 : Reference to Miquel's complaint about the irregulari- 
ties with the dates of Museum Botanicum. Page 334: Reference to the difficulty in choice of 
a successor of Blume. 

1978. Flora Malesiana I, 8: (7)-(16). - Dedication to the memory of F.A.W. Miquel, con- 
taining some notes on Blume. 

& R.S. Cowan. 1976. Taxonomic literature. Ed. 2, vol. 1 : 234-241 (Regnum Vegetabile 94). 

Steenis, C.G.G.J. van. 1941. Natuurwet. Tijdschr. Ned. Ind. 101: 216. - The Planches inedites 
appeared at least before 1883. 

1948. On the date of publication of Blume's Planches inedites. Blumea 6: 263. 

1979. The Rijksherbarium and its contribution to the knowledge of the tropical Asiatic flora. 

Blumea 25: 57-77, especially pp. 60-62. - Blume's endeavours. 

1980. The publication of Blume's Tabellen en Platen voor de Javaansche Orchideeen. 

Miscellaneous Papers Landbouwhogeschool, Wageningen 19: 289-291. 

1986. Blume's Melanges botaniques effectively published, 1855. Taxon 35: 272-273; fac- 
simile of the Melanges: 274-285. 

& M.J. VAN Steenis-Kruseman. 1970. The plates of Javanese plants of Francisco Norona, 

with a revised evaluation of his generic names. In: P. Smit & R.J.Ch.V. ter Laage: Essays in 
biohistory. Regnum Vegetabile 71: 353. - Blume has seen Norona's plates in Java, as well as 



Reinwardt. Incidentally Blume mentioned a few Norona names in the synonymy of his 
Steenis-Kruseman, M.J. van. 1950. Carl Ludwig Blume. Flora Malesiana I, 1: 64-66, 600, 
portr. - Brief personalia; account of Blume's travels and publications. 

1979. Directorate of C.L. Blume. Blumea 25: 35-39. 

Treub, M. 1889. Geschiedenis van 's-Lands Plantentuin te Buitenzorg. Meded. 's Lands Planten- 
tuin 6: 1-79. Batavia. - History of the Botanic Gardens, Bogor, from 1817 till 1844. 

1892. Korte geschiedenis van 's-Lands Plantentuin te Buitenzorg: 7-9, portr. - Short 

history, as above. 

Ule, Witty. Geschichte der Kaiserlichen Leopoldinisch-Carolinischen Akademie der Natur- 
forscher 1852-1882. No. 1071 (not seen). 

Veth, P.J. 1884. Ontdekkers en onderzoekers: 45-149. Leiden. - Mostly on Reinwardt; por- 
trait of 

Vos, C. DE. 1888. Korte schets van de geschiedenis der plantkunde etc.: 91-92. Bolsward. 

Vriese, W.H. de. 1851. Naschrift (to Junghuhn's paper). Ned. Kruidk. Arch. 2: 13-17 (in L). 
- Defending Junghuhn. 

1851. Teregtwijzing van C.L. Blume's naamsverwarring. Alg. Konst-en Letterbode 1850-11: 

35-38. Repr. of 4 pp. in L. - On the reduction of Pinus merkusii to P. finlaysoniana. 

\\ EiGEL, T.O. 1863 (Jan.). Verzeichniss der nachgelassenen Bibliothek von C.L. Blume. Leipzig. 

III-VI + 81 pp. - With portrait of Blume. 
Winkler-Prins, C. 1949. Encyclopaedic ed. 6, 4: 374. 
Wit, H.C.D. de. 1949. 47. Blume. Flora Malesiana I, 4: civ-cv. - A brief account of Blume's 

life; discussion of achievements and main publications. 

1950. History of Malesian botany. 29 pp., unpublished. - Typed copies of letters to and 

from Blume, partly relating to herbarium Reinwardt, but largely official letters on the sale 
and distribution of Flora Javae and Ruinphia. Several derived from the 'Rijksarchief, The 

Zollinger, H. 1841. Tagebuch(ined.), 5 Oct.-31 Dec. - Unpublished diary of Zollinger; typed 
copy by H. Wanner in L. 

Appendix C - References to cited literature 

Arckenhausen, J.C.P. See his biography by H.-G. Griep et al., vide infra. 

Breda, J.G.S. van. 1827-1829. Genera et species Orchidearum et Asclepiadarum quas in 
itinerere per insulam Java collegerunt Dr. H. Kuhl et Dr. J.C. van Hasselt. Ghent. Folio. 15 
fol. & 15 tab. col. 

Da.m, van. 13 Febr. 1832. Ontwerp van eene instructie voor den Directeur van het Rijksherbarium 
(Ministry of the Interior, 5th Div., No. 254 - Concept of an Instruction to the Director of the 

Directions for the director in 14 articles: how to manage the collections, the accommodation, 
the facilities for and availability to other botanists, loans, the making of a catalogue of the col- 
lections, exchange of duplicates, desirability of acquiring collections from civil servants, the fu- 
sion of the University Herbarium with that of the Rijksherbarium, the order that the director 
writes an annual report on the important accessions, and that proposals of the director had to 
go via the Curators of the University. 

A particularly ticklish point was stipulated in art. 10, in which the director was prohibited to 
publish on discoveries of still living persons and explorers without their consent. 
The Instruction was approved by the Minister of the Interior and was stipulated to be effective 
from January 1st, 1831. 


Flora Malesiana 

Griep, H.-G., H. Ullrich &G. Wagenitz. 1977. Johann Christian Arckenhausen (1784-1855). 

In H. Ulrich (ed.), Goslarer Kiinstler und Kunsthandwerker 1: 1-32, illust. (D, 12). 
Hall, H.C. van. 1856. Voorstel omtrent de voortzetting van de uitgave der Flora Javae. In W.H. 

DE Vriese: Tuinbouwflora 3: 365-366. 
Hennipman, E. 1979. The collections of Pteridophytes at the Rijksherbarium. Blumea 25: 

Reinwardt, C.G.C. 1826. Nova plantarum indicarum genera. Syll. Plant. Ratisb. 2: 1-15. 

1 828. Ueber den Charakter der Vegetation auf den Inseln des Indischen Archipels. Ein Vor- 

trag. Kon. Akad. Wiss. Berlin: 1-18. 

Steenis, C. G.G.J. VAN. 1947. Introduction to the Planches inedites Flora Javae (mimeographed). 

- Pamphlet, consisting of a coloured folio plate of Blume's Planches inedites with at the back 

an advertisement for the sale of Blume's works, probably from 1 862 or 1 863 . Copies were sent 

to some selected European libraries. 
& Chew Wee Lek. 1974. Index to C.L. Blume, Museum Botanicum Lugduno-Batavum, 

vol. 2, 1856-1857. Leiden. 24 pp. 
, M.J. VAN Steenis-Kruseman & C.A. Backer. 1954. Louis Auguste Deschamps. Bull. Brit. 

Mus. Nat. Hist., Hist. ser. 1, no 2: 51-68, pi. 13 (a reproduction of the drawing Deschamps 

made of Rafflesia). 
Steenis-Kruseman, M.J. van. 1950. KoUmann's collection of Javan plants. Bull. Jard. Bot. Btzg 

ser. Ill, 18: 463-466. 

1962. Contributions to the history of botany and exploration in Malaysia. 8. Heinrich Burger 

(71806-1858), explorer in Japan and Sumatra. 9. The transfer of the Rijksherbarium from 
Brussels to Holland in 1830. Blumea 11: 495-505; 505-508, 1 photo. 

1979. The collections of the Rijksherbarium. Blumea 25: 29-56. 

Thorbecke, J.R. 11 Nov. 1850. Instructie voor den Directeur van het Rijks-Herbarium te Leijden 
(Ministry of the Interior, 5th Div., No. 254 - Instruction fortheDirector of the Rijksherbarium 
at Leijden). 22 pp. (in L). 

Instruction to replace that of 1832 (see under C: van Dam), consisting of 28 articles. New 
stipulations were: the director should be present on the first three days of the week; not more 
than one family of plants can be borrowed by a single person; the director is prohibited to use 
data from the still living members of the former 'Natuurkundige Commissie' without their per- 
mission; he is not allowed to have a private collection; as to exchange, priority has to be given 
to Dutch botanists and institutes, effective onwards of December 1st, 1850. 
A most peculiar stipulation was in art. 18: anybody could claim to receive duplicates from the 
overseas territories (the names of which had already been printed and the plants described) even 
when nothing was offered in exchange. So it has happened recently that, in cleaning a school 
somewhere in Holland, a set of Javanese sheets was found, obviously claimed by a former en- 
thusiastic teacher who had, it seems, no employ for it. 

Vesque, J. 1883. Catalogue de la Bibliotheque de feu M. J. Decaisne. Avec une notice biogra- 
phique par M. le Dr. Ed. Bornet. Paris. Libraire de la Bibliotheque Nationale: 13. - Listing 
under no 56: 'Blume, Melanges botaniques (Premier et deuxieme numero). Leyde, 1855, br. 
in-8, de 12 pp. -Envoi autogr. de I'auteur a M. Decaisne. 

Vriese, W.H. de. 1858. Reinwardt's Reize naar het Oostelijk gedeelte van den Indischen Archipel 
in het jaar 1821 etc. Amsterdam. 

Appendix D - Notes 

1)- Later it was said that Blume misused the collections and manuscripts of A. Zippelius, a 
gardener of the Botanic Gardens at Buitenzorg (Bogor), who made a long exploration trip to the 
Moluccas, SW. New Guinea, and Timor, where he died. 



Surely Zippelius made a most important collection, but he left no manuscripts at L; we only 
have a box full of old provisional labels. As a matter of fact, P. Bleeker found in the archives 
of the 'Natuurkundige Vereeniging' at Batavia manuscripts and notes of Zippelius that were of- 
fered to Blume about 1850, under the condition that the latter should publish them. Blume never 
replied to this. In fact this request came two decades too late, as Blume had worked on Zippelius's 
material (received through the intermediary of J.B. Spanoghe in ± 1830/31) and published this 
earlier in Rumphia and in the Museum Botanicum. Blume honoured Zippelius by naming the 
Piperaceous genus Zippelia after him. (See also the footnote under D: 4.) 

Blume has also been accused of having left at Bogor no duplicate specimens of the collections 
he took 10 the Netherlands, but this is untrue (see C: van Steenis-Kruseman, 1950; and D: 4). 

As to the Kuhl & van Hasselt collections: they did not add many novelties to what Blume 
himself had collected. The sites where he travelled covered most of theirs, and even far beyond 
eastwards. Besides, the Kuhl & van Hasselt collections came only in Blume's hands in 1828 
when he had already published his Bijdragen (1825-1827) and Enumeratio (1827-1828). And as 
late as 1844 van Breda offered him a packet of notes written by Kuhl and van Hasselt, when 
the main part of Flora Javae (1828-1851) had already been published. 

2) - Ph.F. von Siebold, a most meritorious scientist, withheld his collections from Blume. Most 
of VON Siebold's botanical collections were not made by himself, but by Burger, Textor, Keiske 
and others (see C: van Steenis-Kruseman, 1962). Von Siebold also was a dominating, ambitious 
person. The Flora Japonica was authored by 'Siebold & Zuccarini', but the latter, professor at 
Munich, was the proper author responsible for the research. Von Siebold hardly had any claim 
towards being a botanical taxonomist. As Burger belonged to the 'Natuurkundige Commissie', 
their herbarium should properly go to the Rijksherbarium. Though Burger's share in the under- 
taking was very large - he also wrote a large manuscript on Japanese fishes - von Siebold later 
refused to support Burger's second appointment to the 'Natuurkundige Commissie' for the ex- 
ploration of W. Sumatra, because the latter would not be sufficiently endorsed with scientific 
knowledge (/.c. 501), a most ungracious and unjust gesture. 

VoN Siebold claimed later to have been the saviour of the Rijksherbarium in 1830, whereas his 
sole purpose was to get back specimens collected during his internment by the Japanese in 
Deshima (I.e. 501). Whatever the great merits of von Siebold may have been, these facts throw 
a distinct shadow on his honesty and tolerance regarding other people. 

3) - F. W. Junghuhn was a physician of the army since 1835, but his superior, A.E. Fritze, per- 
mitted him to devote himself to the study of nature. In 1 840 he was charged with making investiga- 
tions in the Batak Lands, W. Sumatra. After his return to Java Junghuhn was appointed a 
member of the 'Natuurkundige Commissie' (1845-1848). Through h\s Reisers dureh 7Gvoand Die 
Battaldnder auf Sumatra it became clear that Junghuhn had amassed a great herbarium, and 
Blume claimed this for the Rijksherbarium. Junghuhn refused, which caused Blume's irritation. 
As Junghuhn was no taxonomist and had made errors in precursory papers (amongst others with 
Lycopodium arboreum), Blume's sharp remarks on this led to a strong mutual animosity between 
him and Junghuhn. 

4) - According to my wife (C: van Steenis-Kruseman, 1950), G.H.J. Kollmann was a German 
senior physician, in the service of the Dutch East Indian army and stationed at Buitenzorg (Bogor) 
in 1821 - 1835, on leave in Europe in 1835-1837. In 1837 he offered the Dutch government a col- 
lection of Javanese plants for sale. ' His letter and material were designated to Blume, who, to 

(I) About the contents of the collection which came in Koilmann's hands more can be found in J. Macii-an, 
Scieniiarum Historia 15 (2), 1973. 1 12 I I.V They comprised zoological collections as well as ethnographical 
ones besides the herbarium specimens. According to Kollmann they were acquired at auctions (presumably 
in Java) and contained not only Blumi collections but also Zippflius plants (M.J. van Steenis-Kruseman). 


Flora Malesiana 

his surprise, found that this was the set of duplicates (more than 4000) of his collection he 
painstakingly left at Buitenzorg when returning to Holland. Kollmann himself never collected. 
Obviously the collection had been stored somewhere in the annexes of the Palace at Buitenzorg, 
adjoining the Botanic Gardens. The curator of the Gardens, James Hooper, was subordinated 
to the Intendant of the Palace. In some way or other Kollmann appropriated this collection. The 
rumour that Blume did not leave duplicates at Buitenzorg appears fully untrue. Why he never 
alluded in print to the curious way in which the Bogor duplicate collection came into his hands, 
can only be guessed at (D: 14). He was either loyal to Kollmann, with whom he had friendly rela- 
tions, or he found it unnecessary to justify himself. Anyway it shows his loyalty to the Buitenzorg 

5) - Both J. Maclean and A. den Ouden (B: 1979) have searched in the 'Rijksarchief, The 
Hague, where all official correspondence by Blume is kept. For a proper biography the period 
1830-1862 should also be covered. Moreover, personal letters will be kept in the archives of 
several botanical institutes as Blume had contacts with many botanists. 

6) - It is quite possible that, as soon as Blume had finished the text for a fascicle of Museum 
Botanicum, he sent it to the printers and assumed it then to be effectively published. In his splen- 
did isolation, surrounded by envious, hostile colleagues and antagonists, Blume did not care 
about their interests. Leiden was at that time a centre where nobody did care about collaboration 
or sympathy, each staff member promoting self-interests; a most unfortunate situation. 

7) - The number of extensive biographies of prominent Dutch botanists is small. I know off-hand 
only those of C.G.C. Reinwardt, Hugo de Vries, W. Beijerinck, F. Junghuhn, J. P. Lotsy, 
F.A.W. MiQUEL, and H.J. Lam. Such biographical studies require much time, and also historical- 
minded people to compose them. If one should like to have a posthumous biography made, it is 
best, in my opinion, to write an autobiography; one ought to think timely of this. 

8) - The diary of H. Zollinger contains notes on his stay at Leiden in 1841, with interesting per- 
sonal information on members of the biological circle at Leiden. Amongst others about the com- 
plaints of Reinwardt that Blume did not give him sufficient honour and published all novelties 
under his own name. But C.A.L.M. Schwaner, a German geologist and member of the 
'Natuurkundige Commissie', said that this was due to the fact that Reinwardt did not publish 
himself, even not his own report on the exploration in East Malesia, and that Reinwardt's 
reasons for not publishing was that he was afraid not to come up to the expectations the botanical 
public had of him. As a matter of fact, the lecture Reinwardt held for this select public, the 'Ver- 
sammlung Deutsche Naturforscher und Aerzte' on 20 September 1828 about the vegetation of 
Malesia, was not exciting, but mediocre (C: Reinwardt, 1828). The same holds for his paper 
Nova plantarum indicarum genera; many genera were assigned to wrong families and several 
others had been described before. Reinwardt's creative efforts lay mainly in the organization 
of botany and cultures in Java, not in research. His report on the exploration of the Moluccas 
was after his death pubhshed in 1858 by W.H. de Vriese (C: 1858), together with a biography. 

Another fact Zollinger mentioned was that it was not due to Blume that P. Korthals aban- 
doned botany. Korthals told Zollinger at the time the first was working out his most important, 
meticulous observations, that botany was an inferior branch of science as compared with 
philosophical and etymological studies, which he found more interesting and scholarly. 

9) - According to Weigel's catalogue (B: Weigel, 1 863), Blume had a very large library, the total 
number of entries being 2123, largely concerning botany (1527 entries). It is peculiar that Blume's 
works are only represented by 9 items. None of his publications on useful and medicinal plants 
were represented. 



10) - As a matter of fact, the majority of biologists, physicians, and explorers in the early part 
of last century concerned with the biology of the Indies were scientists with the German nationali- 
ty or of German descent, e.g., Arckenhausen, Blume, Burger, J.B. Fischer, Hasskarl, 


ScHWANER, VON SiEBOLD, ZiPPELius. Also in South American and African botanical pioneer ex- 
ploration Germans played a prominent role in the former century. 

11) - As to his health, Blume withstood illnesses obviously rather well, probably because he ap- 
plied his own devices, drinking boiled water, etc. He was reported to suffer of fever during his 
trip to Rembang (see A: 1828). In 1826 (Java) Blume complained of illness. In Holland he was 
rather seriously ill about 1829. Early 1850 he suffered of laryngitis. 

12) - H.-C. Griep c.s. (C: 1977) in their biography of J.C.P. Arckenhausen reproduced a letter 
(in the 'Rijksarchief, The Hague) from Blume to the Minister of the Interior at The Hague (d.d. 
27 Dec. 1 832), in which he pleads for the second time for a permanent position of Arckenhausen. 
Blume mentioned that he had 1 500 drawings, mostly from Latour, made in Java. These drawings 
were sketches which should be made ready for reproduction in Flora Javae and often needed to 
be supplemented by details (from herbarium material). Arckenhausen could manage to prepare 
7 or 8 drawings monthly. As the publication of Flora Javae at Brussels needed monthly 12 draw- 
ings for the two instalments, Blume had attracted a certain Mr. Vivien as draughtsman (in 1827) 
and .Mr. Sixtus (in 1828) for keeping pace. Vivien disappeared in 1829 and he was replaced by 
Arckenhausen. The Minister was of the opinion that Arckenhausen should be paid from the 
Flora Javae project funds. The latter worked for Blume at least until 1832, possibly longer. After 
repatriation to Germany Arckenhausen remained draughtsman in Goslar, drawing all kinds of 
plants and animals, mostly for Krebs, Naturgeschichte. After Arckenhausen's death (1855) his 
estate was sold in 1862, among which 134 plates of Flora Javae. In the library of the Natur- 
wissenschaftliche Verein, in volume 19 (portfolio), 190 plates of Blume's work are preserved, of 
which some unfinished sketches. Whether they are originals or printed copies, and whether there 
are unpublished drawings among them, has still to be examined. Plates by Arckenhausen are 
reproduced too in Rumphia, volumes 1-3. 

In Java J.Th. Bik was another artist, originally in the service of Reinwardt, who drew for 

13) - Why the polemic papers between Blume and Junghuhn, de Vriese, and others (see B) 
started as late as 1850 is unclear, because Blume had already in 1844 (see A) reduced Lycopodium 
arboreum - the subject of controversy. Blume's denigrating words accompanying the reduction 
were published by him in Rumphia (3: 219, 221) and these gave offence to Junghuhn and de 
Vriese. Stafleu & Cowan (Taxonomic literature, ed. 2, vol. 1, 1976) gave 1847 as date for this 
part of Rumphia, but it might be that 1849 fits better (as mentioned by Lorentz, c/. Flora Male- 
siana I, 4: clxxii, and also accepted by de Wit). 

14) - Why Blume did not defend himself more openly and publicly is not clear. It is of course 
a fact that one cannot well oppose rumours without published evidence. He was clearly not a very 
militant personality. Blume took action only twice: first, when he revealed the transfer of the 
Rijksherbarium from Brussels and gave honour to Flscher (A: 1831); and second, in defending 
himself against Junghuhn (A: 1850). For the rest he satisfied himself by writing explanatory let- 
ters. Though convinced of his view on the cause of cholera, he did not officially oppose Mulder 
in public. In all these matters I am inclined to believe Blume felt it below his dignity to expose 


Flora Malesiana 

Appendix E - Eponymy 

Blumia C.G.D. Nees 1823, nom. rejic. /^Magnolia L.). 

Blumia K.P.J. Sprengel 1826 ( = Saurauia Willd.). 

Blumea H.G.L. Reichb. 1828 ( ^ \eesta Bll-me). 

Blumea A. P. DC. 1833. 

Blumeodendron Kl'KZ 18"3. 

Blumella van Tieghem 1895 ( ^ Elytranthe Blu )ae ~ Macrosolen Blume). 

Blumeopsis Gacsep . 1920. 

The journal Blumea, official botanical journal of the Rijksherbarium; vol. 1, \934-hodie. 

Epithets for species, blumei, blumii, etc. , are too numerous to enumerate here. 

Appendix F - Honoran distinctions and memberships 

1829 (31 March): Ridder (Knigh:) m de Orde .an de .Nederlandse Leeuw; the Netherlands. 

1851: Legion d'Honneur; France. 

1851: Preussische Rothe .Adler-Ordens, 3. Klasse; Prussia. 

1853: Knight Cross of the Albrechis Order of Sachsen; Saxony. 

1853: Large golden medal for merits from the King of Belgium. 

1822: Council member Bata\iaasch Genootschap van Kunsien en Wetenschappen, Baiavia; Neth- 
erlands Indies. 
1825 (6 Febr.j: Corresponding member of the .Maatschappij van Landbouw en Kruidkunde; the 

182" -.e Koninklijk Instituui van Wetenschappen, Letterkunde en SchooneKunsten 

...- ._... . iiherlands Royal Academy); the .Netherlands. 

1827 (29 June): Member of the Provinciaals Utrechtsch Genootschap voor Kunsten en 
Wetenschappen, Utrecht; the Netherlands. 

1829 (7 Jan.): Member of the Konigliche Botanische Gesellschaft zu Regensburg; Bavaria, Ger- 

1833: Member of the HoUandsche Maatschappij van \\etenschappen, Haarlem; the Netherlands. 

1845: Doctor honoris causa and .Matheseos magister of Leiden University; the Netherlands. 

1851 (7 April): Foreign corresponding member of the Institut de France, Paris; France. 

1853 (May): Ordinary member of the Kaiserliche Akademie fur Naturkunde, .Moscow; Russia. 

1855: Honorary member of the '.Maatschappij ter Bevordering der Geneeskunde', Baden; Ger- 

1855 (31 March): Member of the Koninklijke Akademie van Wetenschappen, Amsterdam; the 

1856 (10 Oct.): Member of the Royal Academy of Sciences, Stockholm; Sweden. 

Member of: 

Caesarea Leopoldino-Carolina Academia Naturae Curiosorum, Bonn; Germany. Cognomen: 

Rumphius secundus. 
Linnean Society of London; England. 

Societas Caesarea Naturae Curiosorum .Mosquensis, Moscow; Russia. 
Societas .Medico-Botanica Londinensis, London; England. 
Natuurkundige Vereeniging van Nederlandsch-Indie, Batavia; Netherlands Indies. 



ace. = according 

Ak. Bis. = Aklan Bisaya (Philip, language) 

Alf. Cel. = Alfurese Celebes (language) 

alt. = altitude 

Anat. = . Anatomy 

.Ap. = .Apayao (Philip, language) 

app. = appendix, appendices 

appr. = approximate 

Apr. = .April 

Arch. = .Archipelago 

atl. = atlas 

auci. div. = auciores diversi; various authors 

auct(t). mat. = auciores malayenses; authors dealing 

with Malesian flora 
auctflj. plur. - auciores plures; several authors 
.Aug. = . August 

Bag. = Bagobo (Philip, language) 
basionym = original name of the type specimen; its 

epithet remains permanently attached to the taxon 

which is typified by it provided it is of the same 

Bg. = Buginese (language) 
Brk. = Bikol (Philip, language) 
Bil. = Bila-an (Philip, language) 
Bill. = Billiton 

Bis. = Bisaya (Philip, language) 
Bon. = Bont6k (Philip, language) 
Born. = Borneo 
Bt = Bukit; mountain 
Bug. = Buginese (language) 
Buic. = Bukidnon (Philip, language) 
c. = circiier, about 

C. Bis. = Cebu Bisaya (Philip, language) 
cf. - confer, compare 
Chab. = Chabecano (Philip, language) 
citations = see references 
cm = centimetre 
c.n. =see comb, now 

comb, now =combinaiio nova; new combination 
CS = cross-section or transversal section of an organ 
c.s. =cum suis; with collaboration 
cum fig. = including the figure 
cur. =curanie: edited by 
D (after a vernacular name) = Dutch 
Daj. = Dyak (language) 
d.b.h. = diameter at breast height 
D.E.I. = Dutch East Indies 
descr. added behind a reference = means that this 

contains a valid description 
diam. = diameter 
Distr. (as an item) = Distribution 
Distr. (with a geographical name) = District 
ditio = the same, see do 
Div. = Division, or Divide 
div. =diversus (masc); various 
do = diiio (Ital.); the same 
Dum. = Dumagal (Philip. language) 
dupl. = duplicate 

E = east (after degrees: eastern longitude) 
E (after a vernacular name) = English 
Ecol. = Ecology 
cd. = edited; edition; editor 
e.g. = exempli gratia; for example 
elah. = elaboravit; revised 
emfend). = emendavit; emended 
em(erg). cd. = emergency edition 
Engl. = English 
etc.. dc. =el cetera; and (the) other things 

ex audi. = ex auciores; according to authors 

excl. =exclusus (masc); excluding, exclusive of 

ex descr. = known to the author only from the de- 
/. (before a plant name) = forma; form 
/. (after a personal name) =filius; the son 

f . (in citations) = figure 

fam. = family 

Feb(r). = February 
fide = according to 

fig. = figure 

f]. =flore, floret (floruit); (with) fiower, flowering 

For. Serv. = Forest Service 

fr. =fructu, fructescit; (with) fruit, fruiting 

Fr. (after a vernacular name) = French 

G. = Gunung (Malay); mountain 

Gad. = Gaddang (Philip, language) 

gen. = genus; genus 

genus delendum = genus to be rejected 

Germ. = German 

geronl. = Old World 

haud = noi, not at all 

holotype = the specimen on which the original de- 
scription was actually based or so designated by 
the original author 

homonym = a name which duplicates the name of an 
earlier described taxon (of the same rank) but 
which is based on a different type species or type 
specimen; all later homonyms are nomenclaturally 
illegitimate, unless conserved 

I. = Island 

ibfid). = ibidem; the same, in the same place 

Ibn. = Ibanag (Philip, language) 

ic. = icon, icones; plate, plates 

ic. inedit. = icon ineditum, icones inedila; inedited 

id. =idem; the same 

i.e. =id est; that is 

If. = Ifugao (Philip, language) 

Ig. = Igorot (Philip, language) 

Ilg. = Ilong6t (Philip, language) 

Ilk. = Iloko (Philip, language) 

in adnol. = in adnotatione; in note, in annotation 

incl. =inclusus (masc); including, inclusive(ly) 

indet. = indetermined 

Indr. = Indragiri (in Central Sumatra) 

inedit. =inediius (masc); inedited 

in herb. = in herbario; in the herbarium 

in lilt. = in lilteris; communicated by letter 

in sched. = in schedula; on a herbarium sheet 

in sice. =in sicco; in a dried state 

in syn. = in synonymis; in synonymy 

Is. = Islands 

Is. (after a vernacular name) = Isinai (Philip, lan- 

Ism. = Isamal (Philip, language) 

isotype = a duplicate of the holotype; in arboreous 
plants isotypes have often been collected from a 
single tree, shrub, or liana from which the holo- 
type was also derived 

Iv. = Ivatan (Philip, language) 

J(av). = Javanese (language) 

Jan. = January 

Jr = Junior 

KIg. = Kalinga (Philip, language) 

Kul. = Kuliman (Philip, language) 

Kuy. = Kuy6non (Philip, language) 

Lamp. = Lampong Districts (in S. Sumatra) 


Flora Malesiana 

Lan. = Lanao (Philip, language) 

lang. = language 

/.c. =loco cilalo; compare reference 

lectotype = the specimen selected a posteriori from 
the authentic elements on which the taxon was 
based when no holotype was designated or when 
the holotype is lost 

livr. = livraison, part 

lice. =l.c. (plur.) 

LS = longitudinal or lengthwise section of an organ 

m = metre 

M = Malay (language) 

Mag. = Magindanao (Philip, language) 

Mak. = Makassar, Macassar (in SW. Celebes) 

Mai. = Malay(an) 

Mai. Pen. = Malay Peninsula 

Mand. = Mandaya (Philip, language) 

Mang. = Mangyan (Philip, language) 

Mar. = March 

Mbo = Man6bo (Philip, language) 

Md. = Madurese (language) 

Minangk. = Minangkabau (a Sumatran language) 

min. part. =pro minore parte; for the smaller part 

mm = milimetre 

Mng. = Mangguangan (Philip, language) 

Morph. = Morphology 

ms(c), MS(S) = manuscript(s) 

Mt(s) = Mount(ains) 

n. = numero; number 

N = North (after degrees: northern latitude); or New 
{e.g. in N. Guinea) 

NE. = northeast 

nee = not 

rieerl. - Netherlands, Netherlands edition 

Neg. = Negrito (Philip, language) 

N.E.I. = Netherlands East Indies 

neotype = the specimen designated to serve as no- 
menclatural type when no authentic specimens 
have existed or when they have been lost; a neotype 
retains its status as the new type as long as no auth- 
entic elements are recovered and as long as it can 
be shown to be satisfactory in accordance with the 
original description or figure of the taxon 

N.G. = New Guinea 

N.I. = Netherlands Indies 

no = numero; number 

nom. =nomen; name (on\y) = nomen nudum 

nom. al. = nomen aliorum; name used by other 

nom. alt(ern). = nomen alternativum; alternative 

nom. cons(erv). = nomen conservandum, nomina 
conservanda; generic name(s) conserved by the In- 
ternational Rules of Botanical Nomenclature 

nom. fam. cons. = nomen familiarum conservan- 
dum; conserved family name 

nom. gen. cons. = see nomen conservandum 

nom. gen. cons. prop. = nomen genericum conser- 
vandum propositum; generic name proposed for 

nom. illeg(it). = nomen illegitimum; illegitimate 

nom. leg(it). = nomen legitimum; legitimate name 

nom. nov. = nomen novum; new name 

nom. nud. = nomen nudum; name published without 
description and without reference to previous pub- 

nom. rejdc.) = nomen rejiciendum; name rejected by 
the International Rules of Botanical Nomenclature 

nom. seminudum = a name which is provided with 
some unessential notes or details which cannot be 
considered to represent a sufficient description 
which is, according to the International Rules of 
Botanical Nomenclature, compulsory for valid 
publication of the name of a taxon 

nom. suhnudum = nomen seminudum 

nom. superfl. = a name superfluous when it was pub- 
lished; in most cases it is a name based on the same 
type as an other earlier specific name 

non followed by author's name and year, not placed 
in parentheses, and put at the end of a citation = 
means that this author has published the same 
name mentioned in the citation independently. 
These names (combinations) are therefore homo- 

Compare 56b line 5-4 from bottom. The same can 
happen with generic names. 

{non followed by abbreviation of author's name) be- 
fore a reference (citation) headed by an other 
author's name = means that the second author has 
misinterpreted the taxon of the first author. 
Compare p. 419a under species 47 the synonym H. 
celebica. Diels misapplied the name H. celebica as 
earlier described by Burck. 

non al. = non aliorum; not of other authors 

non vidi = not seen by the author 

nov. =nova (femin.); new (species, variety, etc.) 

Nov. = November 

n.s. = new series 

n. sp. = nova species; new species 

n. (sp.) prov. = nomen (specificum) provisorium; 
provisional new (specific) name 

n.v. = non vidi; not seen 

NW. = northwest 

Oct. = October 

op.cit. =opere citato; in the work cited 

p. =pagina; page 

P. = Pulau, Pulu (in Malay); Island 

Pal(emb.) = Palembang 

Pamp. = Pampangan (Philip, language) 

Pang. = Pangasinan (Philip, language) 

paratype = a specimen cited with the original descrip- 
tion other than the holotype 

part. alt. = for the other part 

P. Bis. = Panay Bisaya (Philip, language) 

P.I. = Philippine Islands 

pi. = plate 

plurim. = plurimus; most 

p.p. =pro parte; partly 

pr. max. p. =pro maxima parte; for the greater part 

pro = as far as is concerned 

prob. =probabiliter; probably 

prop. = propositus; proposed 

Prov. = Province 

pr.p. =pro parte; partly 

pt = part 

quae est = which is 

9«ooc^ basionym, syn., specimina, etc. =as far as the 
basionym, synonym(s), specimen(s), etc. are con- 

references = see for abbreviations the list in vol. 5, 
pp. cxlv-clxv 

Res. = Residency or Reserve 

resp. = respective(ly) 


Abbreviations and signs 

S = south (after degrees: southern latitude) 

S (after a vernacular name) = Sundanese (language) 

Sbl. = Sambali (Philip, language) 

SE. = southeast 

sec. = secus; according to 

sect. =sectio; section 

sens. ampl. (ampliss.) = sensu amplo (amplissimo); 
in a wider sense, in the widest sense 

sens. lai. =sensu laio; in a wide sense 

sens. sir. (sirictiss.) = sensu siricto (sirictissimo); in 
the narrow sense, in the narrowest sense 

Sept. = September 

seq., seqq. =sequens, sequeniia; the following 

ser. = series 

S.I. =sensu lato; in a wide sense 

S.-L. Bis. = Samar-Leyte Bisaya (Philip, language) 

Sml. = Samal (Philip, language) 

s.n.=sine numero: (specimen) without the collec- 
tor's number 

Sp. = Spanish (language) 

sp(ec). = species; species 

specim. = specimen(s) 

sphalm. = sphalmaie; by error, erroneous 

spp. = species; species (plural) 

Sr = Senior 

5.5. = see sens. sir. 

ssp. = subspecies; subspecies 

s.sir. = see sens. str. 

Stat, now = status nova; proposed in a new rank 

Sub. = Subanum (Philip, language) 

subgfen). = subgenus; subgenus 

subseci. =subsectio; subsection 

subsp. = subspecies; subspecies 

Sul. = Siilu (Philip, language) 

Sum. E.C. = Sumatra East Coast 

Sum. W.C. = Sumatra West Coast 

Suppl. = Supplement 

SW. = southwest 

syn. = synonymum; synonym 

synonyms = the names of taxa which have been re- 
ferred to an earlier described taxon of the same 
rank and with which they have been united on 
taxonomical grounds or which are bound together 

syniypes = the specimens used by the original author 
when no holotype was designed or more specimens 
were simultaneously designated as type 

t. = tabula; plate 

Tag. = Tagalog (Philip, language) 

Tagb. = Tagbaniia (Philip, language) 

Tagk. = Tagaka-6lo (Philip, language) 

Tapan. = Tapanuli (in NW. Sumatra) 

taxon = each entity throughout the hierarchic ranks 
of the plant kingdom which can be described and 
discriminated from other taxa of the same rank 

Taxon. = Taxonomy 

Tg = Tandjung (Malay); cape 

Ting. = Tinggian (Philip, language) 

Tir. = Tirurai (Philip, language) 

transl. = translated 

type = each taxon above the rank of a species is typi- 
fied by a type belonging to a lower rank, for in- 
stance a family by a genus, a genus in its turn by 
a species; a species or infraspecific taxon is typified 
by a specimen. Thenameof ata.xon is nomenclatu- 
rally permanently attached to its type; from this it 
cannot be inferred that the type always represents 
botanically the most typical or average structure 
found in the circumscription of the taxon. 

type specimen = the specimen or other element to 
which the name of a species or infraspecific taxon 
is (nomenclaturally) permanently attached; botan- 
ically a type specimen is a random specimen on 
which the name was based by description. There- 
fore, it does not need to represent the average or 
most typical representative of a population. See 
holotype, isotype, lectotype, syntype, paratype, 
and neotype 

typ. excl. = typo excluso; type excluded 

typ. incl. = typo incluso; type included 

typus = see type and type specimen 

var. = varietas; variety 

var. nov. = varietas nova; new variety 

Vern. = Vernacular 

vide = see 

viz. = videlicet; namely 

vol. = volume 

W = west (after degrees: western longitude) 

Yak. = Yakan (Philip, language) 

± = about 

& = and 

= diameter 

(^ =male (flower, etc.) 

$ = female (flower, etc.) 

5, <^ -bisexual (flower) 

(cJ) (9) = dioecious with unisexual flowers 

(c??) = monoecious with unisexual flowers 

(<? 5) = polygamous 

(??) = polygamous 

oo = many 

> = more than (in size, number, etc.) 

< = less than (size, number, etc.) 
X 2/5 = 2/5 of natural size 

X montana = means that the epithet montana is that 
of a hybrid 



Trees or shrubs (or rarely suffrutices outside Malesia). Leaves simple, alternate, 
often coriaceous, glabrous or with an indumentum on undersurface, margin en- 
tire; petioles often with 2 lateral glands. Stipules 2, minute and caducous to large 
and persistent, usually linear-lanceolate. Inflorescence racemose, paniculate or 
cymose; flowers bracteate and usually bibracteolate; bracts and bracteoles small 
and caducous or larger and enclosing flower or groups of flowers and persistent. 
Flowers actinomorphic to zygomorphic, hermaphrodite or rarely polygamous, 
markedly perigynous. Receptacle campanulate to cylindrical or rarely flattened 
cupuliforum, often gibbous at base; calyx lobes 5, imbricate, often unequal, 
erect or reflexed. Petals 5 (absent in some Neotropical species), inserted on mar- 
gin of disk, commonly unequal, imbricate, deciduous, rarely clawed. Stamens 
indefinite, 2-60 (to 300 in Neotropics), inserted on margin of the disk, in a com- 
plete circle or unilateral, all fertile or some without anthers and often reduced 
to small tooth-like staminodes; filaments filiform, free or ligulately connate, 
short and included to long and far exserted; anthers small, 2-locular, longitudi- 
nally dehiscent, glabrous or rarely pubescent. Ovary basically of three carpels 
but usually with only one developed, the other two aborted or vestigial, various- 
ly attached to (the base, middle or mouth oO receptacle, usually sessile or with 
short gynophore, pubescent or villous; ovary unilocular with two ovules or bi- 
locular with one ovule in each locule. Ovules erect, with micropyle at base 
(epitropous). Style filiform, basally attached; stigma 3-lobed or truncate. Fruit 
a fleshy or dry drupe of varied size, interior often densely hairy; endocarp much 
varied, thick or thin, fibrous or bony, often with a special mechanism for seed- 
ling escape. Seed erect, exalbuminous, the testa membraneous; cotyledons 
amygdaloid, plano-convex, fleshy, sometimes ruminate. Germination hypogeal 
with the first leaves opposite or alternate or epigeal with opposite first leaves. 

An extensive review of the generic limits of the family has been published: G.T. Prance & 
F. White, The genera of Chrysobalanaceae: a study in practical and theoretical taxonomy and 
its relevance to evolutionary biology, Phil. Trans. Roy. Soc. London 320 (1988) 1-184. This con- 
tains full details of taxonomic history, morphology, anatomy, pollen, ecology and distribution 
of the family. A condensed version of these subjects is given here. Details of the Neotropical 
members of the family are given in: G.T. Prance, Chrysobalanaceae, Flora Neotropica 9 (1972) 
1-410. The African members of the family were treated in: F. White, The taxonomy, ecology 
and chorology of African Chrysohalanaceae (excluding Acioa), Bull. Jard. Bot. Nat. Belg. 46 
(1976) 265-350. 

Distribution. Pantropical with 456 species in 17 genera; 365 species in the Neotropics, 57 
in Africa, and 34 in Asia, Malesia and the Pacific. 

Seven genera are native to the Flora Malesiana region and one species of an eighth genus, Chry- 
sobalanus, from Africa and South America, has naturalized in Malesia and Fiji and is therefore 
included in this treatment. All four tribes of Chrysobalanaceae are represented in the region. The 
genera treated here fall into the following tribes of Pranc i- & White: 

Tribe Chrysobalaneae: Chrysobalunus, I.icania, Paraslemon. 

(I) Drawings made by Bobbi Angcll. David Woolcott, KirMcn Tind, and Julia Lokcn; David Johnson 
assisted with ihc distribution maps. 


636 Flora Malesiana [ser. I, vol. lO"* 

Tribe Parinarieae: Hunga, Parinari. 

Tribe Couepieae: Maranthes. 

Tribe Hirtelleae: A tuna, Kostermanthus. 

The genera Atuna, Hunga, Kostermanthus, and Parastemon are confined to the Malesian and 
Pacific region. Licania is predominantly a Neotropical genus (186 species there) with a single 
species in West Africa and three in Malesia. Parinari is a pantropical genus with almost equal rep- 
resentation in all three major regions of the tropics, and Maranthes is predominantly an African 
genus with one abundant and widespread species in Malesia and the Pacific and a single closely 
related species in Central America. 

Morphology. All species of Chrysobalanaceae are woody and most are trees or treelets. All 
are leptocaul. Several, including species of Atuna, Kostermanthus, Licania (Neotropical), Magni- 
stipula (African), Maranthes and Parinari, exceed a height of 30 m and are important constituents 
of the upper forest canopy or are emergents. Six African and Neotropical species belonging to 
Licania, Magnistipula and Parinari are geoxylic suffrutices with massive woody underground 
parts, but rather exiguous aerial shoots which are capable of only limited upward growth and a 
similar form occurs in Parinari nonda in Australia. 

In their architecture and growth-dynamics those Chrysobalanaceae that have been studied ex- 
hibit the model of Troll. This has been demonstrated only in African and Neotropical species. 

Herbarium specimens of Atuna show a distinct pattern of branching which is difficult to 
describe except in terms of development based on the living plant. 

Buttresses are normally absent but frequently well-developed in some species of Parinari and 
Atuna, for example, P. canarioides, P. costata, P. oblongifolia, A. cordata and A. excelsa, and 
the trunk of some species of Parinari, e.g. P. parva and P. gigantea is often fluted at the base. 

The leaves, which are simple and spirally inserted, are frequently arranged distichously. Most 
species have stiff, coriaceous, evergreen leaves which contain abundant silica inclusions. 

Stipules are nearly always present but are sometimes small and caducous. In some Neotropical 
species of Parinari the stipules reach a length of 7 cm, they are up to 4 cm in Parinari parva. In 
Atuna they are prominently keeled, a unique feature in the family. 

The lamina is entire, in all Malesian species. In nearly all species of Parinari, and a few 
Neotropical species of Licania, the veins on the lower surface are extremely prominent and form 
a dense network occupying more than half of the leaf surface so that the stomata are confined 
to relatively small sunken crypts which are densely filled with short curly hairs. 

Foliar glands occur in most, possibly all, species. They secrete nectar which is eaten by ants, 
and function chiefly on young leaves. On mature leaves of herbarium specimens they are not 
always clearly visible. The structure and distribution on the leaf of the glands varies greatly from 
genus to genus and provides characters of considerable taxonomic importance. Small discoid 
glands occur in various places on the lower surface or margins of the lamina in Parastemon. There 
are larger, sometimes ill-defined, glandular areas towards the base of the lamina in Maranthes. 
In Parinari conspicuous glands occur on the petiole. 

The inflorescence is very variable. In Chrysobalanus the few-flowered inflorescence is a short 
raceme of cymules or is cymose throughout, or is a false raceme or a subsessile fascicle. In 
Parastemon the inflorescence is a simple or branched raceme. Hunga and Malesian species of 
Licania have simple or branched racemes of usually congested cymules. More complex mixed in- 
florescences with cymose ultimate units are found in Kostermanthus and Parinari, and the inflo- 
rescence of Maranthes is corymbose. 

Since the inflorescence is usually cymose, at least in part, a distinction between bract and 
bracteole cannot always be drawn. Bracts and bracteoles are usually small but in nearly all species 
of Parinari they are relatively large and enclose small groups of developing flowers. 

In most species the flowers appear to be bisexual, but future field work may show that this is 
not always so. Parastemon urophyllus is said to be polygamodioecious. 

Floral symmetry varies from almost completely actinomorphic, apart from the lateral style, in 

1989] Chrysobalanaceae (Prance) 637 

Chrysobalanus, Parastemon, and most species of Licania to strongly zygomorphic in Kosterman- 
thus. Aciinomorphic flowers are patelliform or shallowly cupuliform, and zygomorphic flowers 
usually have a long receptacle-tube, but in Kostennanthus the strongly zygomorphic flowers have 
a very short receptacle. In the Chrysobalanaceae the receptacle-surface is always lined with nectar- 
secreting tissue, which sometimes, as in Maranthes corymbosa, almost completely fills the tube. 
In most genera the entrance to the receptacle tube is blocked by long straight retrorse hairs, but 
these are lacking in Kostermanthus. In Parastemon the nectariferous lining of the receptacle is 
freely exposed. 

There are always five, completely free, slightly to strongly imbricate sepals which vary from 
subequal in Chrysobalanus to markedly unequal in Kostermanthus. In most genera they are acute 
or subacute but in Kostermanthus and Maranthes they are suborbicular and deeply concave. 

Petals are present in all Malesian species but absent in many Neotropical species of Licania. 
There are always five. They are mostly caducous. In shape they vary from linear-spathulate 
(Chrysobalanus) to orbicular. They are usually subequal, but in Kostermanthus they are very un- 
equal in shape and size and are strongly unguiculate. 

Stamens vary in number from two in Parastemon urophyllus to 40 in Maranthes. In Chrysoba- 
lanus, most species of Licania, Parastemon versteeghii, and Maranthes they form a complete or 
almost complete circle round the entrance to the flower and all or most are fertile. Otherwise the 
fertile stamens are inserted unilaterally opposite the carpel. Staminodes are frequently present op- 
posite the style. In several genera the filaments appear to be united at the base, but it is sometimes 
difficult to decide whether this represents true union or whether the filaments are free but inserted 
on a development of a receptacular rim. In Maranthes the stamens are inserted in two or more 
rows on the outer surface of what appears to be a receptacular annulus. In length the filaments 
vary from much shorter than the calyx, as in Hunga, Parastemon and some species of Licania, 
to very much longer in Maranthes. In Kostermanthus the filaments are united for at least half of 
their length to form a conspicuous ligule. 

The gynoecium fundamentally is composed of three carpels which are free except for the 
gynobasic style. In most species there is only one functional carpel, though one or two small 
rudimentary carpels can sometimes be seen. Due to the development of a false dissepiment the 
ovary is bilocular in Hunga, Parinari, and A tuna. 

The fruit is basically a drupe but there is considerable variation in detail, apparently associated 
with dispersal and germination. In Chrysobalanus, Parastemon and Hunga the endocarp has a 
smooth surface and is sharply differentiated from the mesocarp. In the other genera the differen- 
tiation is less well-defined. In Chrysobalanus and Hunga, seedling escape is effected by means of 
longitudinal lines of weakness. In Parastemon and Maranthes two large lateral plates fall away 
permitting the seedling to emerge. In Parinari there are two small basal 'plugs' or obturators. All 
other genera seem to lack specialized means of seedlings escape. 

In Chrysobalanus, Licania, Parastemon, Parinari, and A tuna, germination is cryptocotylar, 
whereas in Maranthes it is phanerocotylar. 

Vegetative Anatomy. - Leaf anatomy. Indumentum, if present, consisting of long unicel- 
lular hairs. Variously positioned glands (cxtrafloral nectaries) with slender upright epidermal 
secretory cells commonly present. Wax present as platelets (Fhhrlnbach &, 1988). 
Siomata mostly paracytic, confined to the lower leaf surface. Upper epidermis often composed 
of tall cells; with mucilaginous inner walls in some species. Hypodermis often present. Mesophyll 
entirely composed of palisade-like cells, more rarely dorsivcntral and differentiated into palisade 
and spongy tissue. Astcrosclcrcids occasionally present in mesophyll. Veins mostly with scleren- 
chyma sheaths including sclercids with U-shaped wall thickenings, sometimes vertically Iranscur- 
rent. Midrib and distal end of petiole with a closed vascular cylinder, with or without additional 
adaxial or medullary collateral bundles. Silica bodies and silicified cell walls common, especially 
in epidermis. 

Young stem. Cork arising superficially. Pcricyclic sclercnchyma ring composed of fibres and 

638 Flora Malesiana [ser. I, vol. 10"^ 

sclereids with U-shaped wall thickenings. Secondary phloem occasionally with secretory (tannin?) 
cells. Sieve tube plastids of the S-type (Behnke, 1984). Silica bodies often present in pericycle, 
phloem and xylem rays, and in pith. 

IVood anatomy. Growth rings absent or, if present, defined by differences in the spacing of 
tangential parenchyma bands. Vessels diffuse, often in a weakly oblique pattern, (almost) exclu- 
sively solitary, tending to be of two distinct sizes, the larger ones very wide (200-300 [im). Vessel 
perforations simple. Tyloses often present in heartwood, sclerotic in some species. Vessel-ray pit- 
ting including elongate horizontal or oblique to almost vertical pits with strongly reduced borders, 
often unilaterally compound. Fibres often thick-walled, with distinctly bordered pits throughout 
the tangential walls, and in the radial walls often confined to fibre-ray contacts (fibre-tracheids); 
in contact with vessels often less thick-walled and with biseriate bordered pits ( == vasicentric 
tracheids). Parenchyma in fine uniseriate or locally bi(-tri)-seriate, regular or irregular wavy tan- 
gential bands. Parenchyma strands typically long, of up to 16 cells. Some axial parenchyma cells 
with spiral thickenings in Atuna p.p., Licania, Maranthes p.p., and Kostermanthus (Jer Welle, 
1975). Rays predominantly uniseriate, but in some taxa also biseriate, typically weakly heteroge- 
neous with (often weakly) procumbent central cells and one row of square to upright marginal 
cells (Kribs type III), sometimes homogeneous and composed of procumbent cells only. Silica 
bodies universally present in ray cells, more rarely in axial parenchyma cells. Rhomboidal crystals 
in chambered axial parenchyma cells noted in Parastemon. 

Taxonomic notes based on vegetative anatomy. The above general anatomical description is 
based on the literature (for leaf and young stem anatomy mainly Kuster, 1897, as abstracted by 
SoLEREDER, 1899; and Prance, 1972, and Prance & White, 1988, for wood anatomy from many 
sources), amplified with original observations on slides present in the Rijksherbarium at Leiden. 
A number of anatomical characters may prove to be of considerable taxonomic significance at 
the genus or species level (mucilaginous leaf epidermis, distribution of silica grains in leaves, 
young stem, and wood, vascular pattern and sclerenchyma support of leaf veins and petiole, fibre 
and sclereid distribution pattern of the mature bark (Roth, 1981), spiral thickenings in axial 
parenchyma cells of the wood, ray width and histology, etc.). However, for the Malesian Chryso- 
balanaceae their diagnostic value remains largely untested. On the whole the Chrysobalanaceae 
are anatomically rather homogeneous, and as repeatedly emphasized, quite distinct from the 
Rosaceae. Anatomically Chrysobalanaceae are also distinct from the numerous families to which 
they have been compared in the search for closest relatives. 

References: Behnke, Ann. Missouri Bot. Gard. 71 (1984) 824-831; Desch, Manual of 
Malayan Timbers 2 (1954) 474-485; Burgess, Timbers of Sabah (1966) 434-436; Fehrenbach 
& Barthlott, Bot. Jahrb. 109 (1988) 407-428; Furuno, Anatomy of Papua New Guinea Wood 
(Continued), Res. Report of Foreign Wood 8, ShimaneUniv. (1979); Hayashi c.5., Micrographic 
Atlas of Southeast Asian Timber, Kyoto Univ. (1973); Lecomte, Les bois de I'lndochine (1926) 
59-61; Metcalfe & Chalk, Anatomy of the Dicotyledons 1 (1950) 550-553; Moll & 
Janssonius, Mikrographiedes Holzes der auf Java vorkommenden Baumarten 3 (1914)222-230; 
Prance, Flora Neotropica 9 (1972) 1-19; Prance & White, Phil. Trans. Roy. Soc. Lond. B 320 
(1988) 1-184; Roth, Encycl. Plant Anatomy 9, 3 (1981) 286-295, 402-403; Solereder, 
Systematische Anatomic der Dicotyledonen (1899) 341-351; Ter Welle, Acta Bot. Neerl. 24 
(1975) 397-405; lAWA Bulletin 1976/2 (1976) 19-29; Ter Welle & Detienne, Flora of the 
Guianas A 85 (1986) 109-126. - P. Baas. 

Palynology. The pollen of Chrysobalanaceae is very uniform, but is different from that of 
Rosaceae. It is of little value for distinguishing between the genera of Chrysobalanaceae or for 
arranging them in groups. 

Most species have grains with three furrows, but some species have three or four; there are no 
special features except occasional equatorial constrictions. With light microscopy the pores are 
indistinct, and in some species are difficult to observe. The grains are usually distinctly triangular 
in shape in polar view, except when four-furrowed; they are elliptical to circular in equatorial view 

1989] Chrysobalanaceae (Prance) 639 

and are oblate-spheroidal, prolate-spheroidal or subprolate in shape as indicated by the ratio: 
polar length x 100, divided by the equatorial length = 85-150. The size is very variable from one 
genus to another; the polar area is usually small, sometimes medium, but never large. The exine 
is medium to rather thick with very little patterning; it is usually scabrous to verrucose, but never 

The pollen of Chrysobalanaceae and Rosaceae is similar but readily distinguishable. The 
former is markedly triangular in polar view in the expanded grain, whereas in Rosaceae it is never 
more than weakly triangular. Most Rosaceae have more distinctive pores, and many have more 
patterning on the wall. A feature that occurs frequently in the Rosaceae is a distinct wedge-shaped 
protrusion from the middle of the furrow, obvious in polar view, which does not occur in Chryso- 

Erdtman (1952) states 'pollen morphological objections cannot be raised against regarding the 
Chrysobalanaceae diS a separate family.' Our own study o{ Rosaceae X)o\\tr\ {sensu lato) confirmed 
that three main types of pollen occur: the Rosaceae sensu stricto, the Chrysobalanaceae , and the 
Seuradoideae types (Prance, 1963). The differences between pollen of Chrysobalanaceae and 
Rosaceae are, however, comparatively small. By contrast, the pollen of the Tropaeolaceae, 
Geraniaceae, Limnanthaceae, Linaceae, Polygalaceae, and Sapindaceae, families which various 
phylogenists (Hallier, 1923; Bonne, 1926; Hauman, 1951; Gutzwiller, 1961) have suggested 
are closely related to Chrysobalanaceae, is very different. Pollen morphology thus provides 
reasons for keeping the Chrysobalanaceae near to the Rosaceae in the Rosales, and not for remov- 
ing it to the Geraniales or Sapindales. 

The pollen of Chrysobalanaceae is so uniform that it does not provide good generic characters. 
Kostermanthus heteropetala is distinct from all other Chrysobalanaceae examined, including 
Daciyladenia (Africa) and Acioa (America) with which it shares a staminal ligule, in having three 
swellings on each of the triangular sides of the grain in polar view. Apart from Kostermanthus 
no other genus is clearly definable on pollen characters. 

References: Bonne, C. R. Hebd. Seanc. Acad. Sci. Paris 182 (1926) 1404-1406; Erdtman, 
Pollen morphology and plant taxonomy, Angiosperms (1952) 380-383; Gutzwiller, Bot. Jahrb. 
81 (1961) 1-49; Hallier, Beih. Bot. Centralbl. 39(1923) 1-178; Hauman, Bull. Jard. Bot. Etat 
Brux. 21 (1951) 167-198; Prance, A taxonomic study of the Chrysobalanaceae. Thesis, Oxford 

Phytochemistry. Chemical knowledge about the family Chrysobalanaceae is still scanty. 
Hegnauer (1973) treated it as Chrysobalanoideae sub Rosaceae. Chrysobalanaceae are note- 
worthy for their tendency to accumulate silica (SiOz) in leaves and in the wood where usually every 
ray cell contains one globular silica inclusion. Leaf flavonoid patterns are dominated by the 
flavonols quercetin and kaempferol; some taxa also have myricetin. Proanthocyanidins (formerly 
called leucoanthocyanidins), i.e. condensed tannins, were demonstrated to be present in leaves of 
few species of Chrysobalanus, Licania, and Parinari, but galli- and ellagitannins have not yet been 
traced in the family. The recent flavonoid investigation of 21 species of f*ormcrr/(CoRADiN, Gian- 
NASi & Prance, 1985) resulted in the identification of a number of 3-glycosides of kaempferol, 
quercetin and myricetin, and showed restriction of myricetin glycosides to four African species; 
dihydroquercetin('taxifolin')-3-glycosides were noticeable only in Asian Parinari insularum from 
the Pacific islands and vicenin-like C-glycoflavones only in a few African populations of P. ex- 
celsa. Myricetin was also observed in leaves of Licania macrophylla which besides has much con- 
densed tannins in all parts, saponins in leaf, pericarp, seed, and stem and root bark; alkaloids 
in stem and root bark (Grenand, Morhtti & Jacquhmin, 1987). Cyanogenic glycosides which 
are characteristic of a number of Rosaceous taxa have not been traced in Chrysobalanaceae 
hitherto. The most noteworthy chemical character known from the family at present is the fatty 
acid pattern of their seed triglycerides; conjugated iricnoic and tetraenoic CiK-acids such as alpha- 
claeostearic and parinaric acids are present as major fatty acids in seed oils of species of Chrysoba- 
lanus, Licania, and Parinari s. I. (i.e. including Atuna, Maranthes and the African Neocarya). 

640 Flora Malesiana [ser. I, vol. 10'* 

This character, however, which Hnks Chrysobalanaceae biochemically with Prunoideae (same 
type of seed oils in some Prunus s.l. species) seems not to be universal in the family. According 
to Jones & Earle (1966) seed kernels of a species of Couepia (Central & South America) con- 
tained an oil without conjugated unsaturation. Still too Httle is known from the chemistry of this 
taxon to allow a sound chemotaxonomic discussion. 

References: Coradin, Giannasi & Prance, Brittonia 37 (1985) 169-178; Grenand, Moretti 
& Jacquemin, Pharmacopees traditionelles en Guyane, ed. Orstom, Paris (1987); Hegnauer, 
Chemotaxonomic der Pflanzen 6 (1973) 84-130; Jones & Earle, Econ. Bot. 20 (1966) 137; 
Prance & White, Phil. Trans. Roy. Soc. London B 320 (1988) 28-29. - R. Hegnauer. 

Dispersal. The fruits of Chrysobalanaceae are very uniform in basic structure but remark- 
ably diverse in functional detail. Despite their uniformity they have become adapted to a wide 
range of dispersal agents, sometimes within a single genus or species; however, few species have 
been studied in the field. 

Chrysobalanus icaco ssp. icaco is dispersed by ocean currents, and also by bats, rodents and 
monkeys, and possibly by birds; C. cuspidatus is said to be dispersed by birds. 

Some Neotropical species of Licania are bat-dispersed, whereas the fruits of several South 
American riverine species float and are also eaten by fish; those of the African species L. elaeo- 
sperma are also transported by water. The Malesian species L. splendens is dispersed by the fruit 
pigeon Ducula aenea. 

Various species of Parinari are known to be dispersed by bats, elephants, baboons and other 
primates, a scatter-hoarding squirrel, fruit pigeons, rheas, emus, agoutis and fish. Species of 
Couepia, Licania and Parinari are frequently eaten by bats in the Neotropics. 

Maranthes corymbosa is dispersed by birds, most notable hornbills and fruit pigeons, and, at 
least for short distances, by a scatter-hoarding squirrel. The fruits of some African species are 
eaten by monkeys which are possibly mainly destructive. 

Atuna is dispersed by ocean currents and a scatter-hoarding squirrel and possibly by wild pigs. 

Uses. Members of the Chrysobalanaceae are used by the local people everywhere, for 
building, fuel, charcoal and in folk medicine. The fruits and seeds of some species are highly 
esteemed, and others are eaten in times of scarcity; some are used in the preparation of alcohoUc 
beverages. At present, Chrysobalanaceae are only of local importance commercially, but, with 
improved communications and technology, their potential as a source of construction timber, 
fruits, and edible and industrial oils appears to be promising. 

The Malesian standard timber name for various genera of Chrysobalanaceae is merbatu. 

Edible fruits and seeds. Chrysobalanus icaco is tinned and bottled in syrup and sold in Colom- 
bia and Venezuela under the name Icacos. The fruit of several Neotropical species of Couepia and 
Parinari are eaten. In Amboina a dish called Koku koku is prepared from the mashed seeds of 
Atuna excelsa mixed with raw or fried small fish, ginger, onions, chillies and lime juice. 

Wood. Despite the large supplies of Chrysobalanaceae wood potentially available, commercial 
sawn timber is produced only in relatively small amounts. This is because its high silica content 
blunts even tungsten-tipped saws. Because the wood of many species is resistant to marine borers, 
it is used throughout the tropics for piers and other marine constructions. 

Caulking and waterproofing agent. In the Solomon Islands the principal use oi Atuna excelsa 
sensu lato is for caulking the seams of plank-built canoes. The seeds, which are known as 'putty 
nut' are pounded to a putty-like consistency. After application the putty hardens and darkens, 
but if exposed too long to the sun it cracks, so canoes drawn up on the beach are often kept in 
the shade of sheds. In the central and south-eastern Solomons it is used for setting shell inlay in 
wood bowls, figures and other articles. The north-western Solomon Islanders also use it for water- 
proofing bottles made from gourds. In the Admiralty Islands (Manus) coiled baskets are coated 
with it to make them waterproof (B.A.L. Cranstone, in litt., 14 June 1983). 

History oi Parinari. The taxonomic history of Parinari is complex. At least some species 
of all Malesian genera except Chrysobalanus, and Parastemon have at one time or another been 
placed in Parinari. 

1989] Chrysobalanaceae (Prance) 641 

All species of Aiuna and Maranthes have been included in Parinari. Despite their considerable 
differences from Parinari sensu stricto in virtually all other respects, these genera have one feature 
in common - a bilocular ovary. It was the adoption of this character as a generic criterion, 
especially by Bentham (1849), that led to the increasingly artificial nature oi Parinari. As Parinari 
became more and more heterogeneous even some species with unilocular ovaries were included, 
for example, the species now placed in Kostermanthus. 

In the original description of Parinari, which was based on P. campestris and P. montana from 
French Guiana, Aublet (1775) mentioned the bilocular ovary, but he does not appear to have at- 
tached much importance to it. 

De Jussieu (1789), who brought all previously described genera of Chrysobalanaceae together 
for the first time, knew some of them only from the original descriptions and illustrations. His 
implication that Parinari differs from the other genera principally in its bilocular ovary seems to 
have laid the foundations for the subsequent confused history of the group. 

De Jussieu was the first to extend the concept of Parinari to another continent by citing in 
synonymy two manuscript names of Adanson from Senegal, Mampata and Neou. The former 
was subsequently described as P. excelsa and the latter as P. macrophylla by Sabine. 

The following year, in his Prodromus, De Candolle (1825), who only knew the four species 
mentioned above, divided Parinari into two sections. Section Petrocarya (correctly section 
Parinari) was based on a superfluous generic name which Schreber (1789) substituted for the 
earlier Parinari. It included Aublet's original species. Section Neocarya was based on P. 
senegalensis DC. [now Neocarya macrophylla (Sabine) Prance], but P. excelsa was associated 
with it, probably because its type-description is inadequate to characterize it properly. Parinari 
macrophylla is not mentioned by De Candolle. He was also apparently unaware of the first true 
Parinari to be described from Asia, P. sumatrana Benth., which had been described by Jack in 
the illegitimate genus Petrocarya in 1822. De Candolle indirectly emphasized the importance of 
the bilocular ovary of Parinari by describing the ovary of all other genera as unilocular. 

During the first half of the nineteenth century, in addition to Neocarya macrophylla, a few 
other species, which belong to other genera, were described in Parinari or its illegitimate synonym 
Petrocarya, because of their bilocular ovary. Thus Jack ( 1 822) described Petrocarya excelsa (now 
Atuna excelsa), and Bentham (1840) published Parinari coriacea (now Exellodendron coriacea), 
but it was Bentham's treatment of Parinari in Hooker's Niger Flora (1849) that firmly establish- 
ed Parinari as an artificial genus. 

Whereas earlier workers had implied that the bilocular ovary is a diagnostic character of 
Parinari, Bentham referred to the spurious dissepiment which separates the ovules as 'the essen- 
tial character.' Bentham divided Parinari into three sections as follows: 

Section 1: Petrocarya (correctly Parinari) included the African species P. excelsa and P. 
curatellifolia, all the known American species including P. coriacea (now Exellodendron cor- 
iaceum), and, with some doubt, three species Bentham had not seen himself, namely P. 
sumatrana Benth. (a true Parinari), P. glaberrima Hassk. (now Atuna excelsa) and P. scabra 
Hassk. (now Atuna scabra). 

Section 2: Sarcostegia Benth. included two new species, P. polyandra (now Maranthes polyan- 
dra) and P. griffiihiana (now Maranthes corymbosa), and, with some doubt, also P. jackiana 
Benth. (based on Petrocarya excelsa, now Atuna excelsa) which Bentham had not examined. 

Section 3: Neocarya DC. contained P. macrophylla (now Neocarya macrophylla) and its 
synonym P. senegalensis. 

Bentham's circumscription of Parinari was probably much wider than he imagined, largely 
because of the inclusion of the Asian species he only knew from the literature. He appears to have 
adopted it with some reservation. Parinari polyandra has c. 40 fertile stamens and Bentham men- 
tions that this, in conjunction with the glandular leaves and fleshy 'calyx', might 'suggest the es- 
tablishment of a distinct genus.' He clearly believed that the stamen number of Parinari varies 
more or less continuously, but the evidence he cites is partly on the species he had not studied. 

642 Flora Malesiana [ser. I, vol. 10"^ 

Bentham's circumscription of Parinari included five genera which are now regarded as distinct, 
namely, in additon to Parinari itself, Atuna Rafin., Exellodendron Prance, Maranthes Blume 
and Neocarya Prance. Two of these from Malesia had enjoyed a brief period of generic recogni- 
tion. Thus, Maranthes was described by Blume in 1825, but three years later he transferred the 
type species to his illegitimate Exitelea. A tuna was described by Rafinesque in 1 838, but remained 
disregarded for more than 100 years, though one of its species was independently described by 
Hasskarl in 1842 as the type of his new genus Cyclandrophora. It appears that Hasskarl had 
little faith in his new genus for he united it with Parinari within a year of its publication, although 
it has little in common with the latter, other than the bilocular ovary. 

Since Bentham (1849) nearly all species of Chrysobalanaceae with false dissepiment (and even 
some without) were automatically placed in Parinari regardless of any other consideration. 

As new species now placed in Exellodendron, Maranthes and Atuna were described they were 
all placed in Parinari. Likewise, equally disparate elements which are now placed in Bafodeya 
Prance, Hunga Pancher ex Prance and Kostermanthus Prance joined the assemblage. 


based on flowering material 

1. Stamens free, not united into a ligule; petals not clawed, ovary uni- or bilocular. 
2. Ovary unilocular, inserted at or near base of receptacle. 
3. Inflorescence a panicle of cymules; fertile stamens 7-26. 
4. 'Stamens 15-26, the filaments hairy, exserted 1. Chrysobalanus 

4. Stamens 7-10, the filaments glabrous, included 2. Licania 

3. Inflorescence racemose; fertile stamens 2 or 5 3. Parastemon 

2. Ovary bilocular, inserted at mouth or midway up receptacle. 

5. Fertile stamens 6-8(-9), markedly unilateral, the filaments equal or not exceeding the calyx lobes. 
6. Lower leaf surface glabrous or lanate, with stomatal cavities; bracteoles not enclosing small groups of 
flowers; inflorescence a panicle of cymules; ovary inserted midway up receptacle 4. Hunga 

6. Lower leaf surface usually areolate with stomatal cavities; bracteoles enclosing small groups of flowers; 
inflorescence a much-branched panicle; ovary inserted at mouth of receptacle 5. Parinari 

5. Fertile stamens 10-50, usually inserted around complete circle; the filaments far exserted beyond calyx 

7. Stamens 10-25; inflorescence little branched panicles, or racemes 6. Atuna 

7. Stamens c. 45; inflorescence much-branched corymbose panicles 7. Maranthes 

1. Stamens united into a strap-shaped ligule; the 2 anterior petals unguiculate and enveloping the ligule in 
buds; ovary unilocular 8. Kostermanthus 


based on fruiting material 

1 . Epicarp crustaceous-verrucose; mesocarp thick, hard, fibrous; endocarp breaking up irregularly on germi- 
nation; cotyledons at least slightly ruminate. 
2. Stamens free to base (can often be seen persistent around base of young fruit). Cotyledons ruminate 

6. Atuna 

2. Stamens united into a unilateral ligule. Cotyledons only slightly ruminate 8. Kostermanthus 

1. Epicarp smooth and glabrous or distinctly lenticellate but not crustaceous; if lenticellate then endocarp 

opening by a pair of basal stoppers to allow seedling escape. Cotyledons not ruminate. 

3. Epicarp lenticellate; opening by a pair of basal stoppers to allow seedling escape, always thick and woody; 
fruit bilocular, but often only one loculus developing seed 5. Parinari 

3. Epicarp glabrous and smooth without lenticels; opening by lateral plates, longitudinal lines or no special 
mechanism of seedling escape; fruit uni- or bilocular. 

4. Endocarp opening by a pair of lateral plates to allow seedling to escape; endocarp thick and woody or 
thin and bony. 

5. Fruit unilocular; endocarp very thin, bony 3. Parastemon 

5. Fruit bilocular; endocarp thick, woody 7. Maranthes 

1989] Chrysobalanaceae (Prance) 643 

4. Endocarp not opening by lateral plates, usually opening longitudinally; endocarp thin and bony. 
6. Fruit usually bilocular, 1.5-5 cm long, sometimes sagittate with a distinct stipe, not ridged 

4. Hunga 
6. Fruit unilocular, either 1-1.3 cm long, ellipsoid or 2-5 cm long and ridged. 

7. Fruit ridged. Leaves orbicular 1. Chnsobalanus 

7. Fruit smooth, not ridged. Leaves oblong to elliptic 2. Licania 


Linn. Sp. PI. 1 (1753) 513; DC. Prod. 2 (1825) 525; Hook./, in Benth. & 
Hook./., Gen. PI. 1 (1865) 606; Hook./ in Mart., Fl. Bras. 14 (2) (1867) 7; 
Prance, Fl. Neotrop. 9 (1972) 14. - Fig. 1. 

Shrubs or small trees. Stipules small, connate-axillary, caducous. Leaves 
glabrous on both surfaces, without stomatal crypts. Petioles eglandular. Inflo- 
rescence terminal or a.xillary cymules or a panicle of cymules. Bracts and brac- 
teoles to 2 mm long, eglandular, not enclosing groups of flower buds. Flowers 
hermaphrodite. Receptacle campanulate, symmetric, tomentose on exterior and 
interior; calyx lobes 5, acute, equal. Petals 5, longer than calyx lobes, not 
clawed. Stamens 15-26, all fertile, inserted on margin of disk; filaments hairy, 
united in groups for half length, exserted. Ovary inserted at base of receptacle, 
densely pilose; carpel unilocular, with 2 ovules. Style pubescent. Fruit a small 
fleshy drupe, epicarp smooth and ridged, endocarp hard, thin, glabrous on in- 
terior, with 4-8 prominent longitudinal ridges which correspond to lines of 
fracture for seedling escape. 

D i s t r . Three species, one in West Africa and the Neotropics, two confined to the Neotropics. One species 
naturalized in Malesia and Fiji. 
Uses. Edible fruit. The shrub is used for the stabilization of dunes. 

1. Chr>$obalanus icaco Linn. Sp. PI. 1 (1753) 513; rical, tomentose on exterior and interior. Calyx lobes 

Browne, Nat. Hist. Jamaica (1756) 250; Jacq. Sel. rounded to acute, tomentellous on both surfaces. 

Stirp. Am. Hist. (1763) 155; DC. Prod. 2(1825)525; Pe/o/s white, glabrous, exserted. S/flrne'n5 15-26, the 

Hook./, in Mart., Fl. Bras. 14(2) (1867) 7; Prance, filaments joined for up to half of length in small 

Fl. Neotrop. 9 (1972) 15; Smith, Fl. Vit. Nov. 3 groups, densely hairy, exserted. Ovary at base of 

(1985) 50. - Fig. 1. receptacle, pilose. Fruit ovate to obovate, 2-5 cm 

Shrub or small tree to 5 m tall, the branches gla- long; epicarp smooth with longitudinal ridges; 

brous and lenticellate. S//pM/« 1-3 mm long, cadu- mesocarp thin and fleshy; endocarp thin, hard, 

cous. Leaves orbicular to ovate-elliptic, 2-8 by 2-6 ridged on exterior. 

cm, retuse, rounded or with short blunt acumen at Distr. Neotropics, mainly in coastal areas; West 

apex, subcuneate at base, glabrous on both surfaces; & Central Africa, naturalized in Fiji, cultivated in 

petioles 2-4 mm. Inflorescences small terminal and Vietnam; in Malesia cultivated in Singapore where it 

axillary cymules or panicles of cymules, the rachis has escaped and naturalized. Fig. ID. 

and branches grey-puberulous. Flowers 4-6 mm Ecol. Dunes, beaches and coastal scrub, 

long. /?ecep/flc/^ campanulatc-cupuliform, symmct- Uses. Edible fruit. 


Flora Malesiana 

[ser. I, vol. 10^ 


1 cm 

Fig. 1. Chrysobalanus icaco Linn. A. Detail of flower; B. habit; C. fruit; D. distribution in Malesia. 


Chrysobalanaceae (Prance) 


Fig. 2. Licania splendens (KoRTH.) Prance. A. Habit, xO.5; B. flower, x9; C. flower section, x9; D. fruit, 
X 1 (A-C Elmer 20916, D Ramos & Convocar 83437). 


AuBL. Hist. PI. Guiane Fr. 1 (1775) 1 19, t. 45; DC. Prod. 2 (1825) 527; Hook. 
/. in Benth. & Hook./. Gen. PI. 1 (1865) 606; Fritsch, Ann. Naturh. Mus. Wien 
4 (1889) 33; Focke in E. & P. Nat. Pn. Fam. 3, 3 (1891) 58; Prance, F1. 
Neotrop. 9 (1972) 21; Prance & Whitm. Tree Fl. Malaya 2 (1973) 328; White, 
Bull. Jard. Bot. Nat. Belg. 46 (1976) 280; Prance, Brittonia 31 (1979) 94. - 
Moquilea Aubl. Hist. PI. Guiane Fr. 1 (1775) 521, t. 208; DC. Prod. 2 (1825) 
526; Hook./, in Benth. & Hook./, Gen. PI. 1 (1865) 606; Focke in E. & P. 
Nat. PH. Fam. 3, 3 (1891) 58. - Dahuronia Scop. Introd. (1777) 217, nom. illeg. 
- Hedycrea Schreb. in Linn., Gen. PI. ed. 8, 1 (1789) 160, nom. illeg. - 
Angelesia Korth. Ned. Kruidk. Arch. 3 (1854) 384; Boerl. Handl. Fl. Ned. 
Ind. 1 (1890) 424; Burk. Diet. (1935) 159; Corner, Wayside Trees (1940) 526; 
Hutch. Gen. Flow. PI. 1 (1964) 191. - Trichocarya Miq. Fl. Ind. Bat. 1, 1 
(1855) 358; ibid. 6 (1858) 1084, p.p. quoad T. splendens tantum. - Geobalanus 
Small, Fl. Miami (1913) 80; Hutch. Gen. Flow. PI. 1 (1964) 191. - Coccomelia 
Ridley, J. Str. Br. Roy. As. Soc. n. 82 (1920) 183; Fl. Mal. Pen. 1 (1922) 
671. - AfrolicaniaMiLDBK. Notizbl. Bot. Gart. Bcrlin-Dahlem 8 (1921) 483. - 
Fig. 2. 

Small to large trees. Stipules small, free, caducous. L^crv^^ glabrous on both 
surfaces, without stomatal crypts. Petioles eglandular. Inflorescence a panicle 
of cymules. Bracts and bracteoles to 1.5 mm long, membraneous, eglandular, 
not enclosing groups of flower buds. Flowers hermaphrodite. Receptacle cam- 
panulate, slightly asymmetric, tomcntose on exterior, tomentosc within; calyx 


Flora Malesiana 

[ser. I, vol. 10^ 

lobes 5, acute, unequal. Petals 5, small, not exceeding the calyx lobes, not 
clawed. Stamens 7-10, all fertile, inserted on margin of disk; filaments gla- 
brous, included, slightly united at base. Ovary inserted at or near base of recep- 
tacle, pilose on exterior; carpel unilocular, with 2 ovules. Style pubescent at 
base, the stigma capitate. Fruit a small, fleshy drupe, narrowed to a shortly stip- 
itate base; epicarp smooth, not ridged, glabrous, not lenticellate; mesocarp thin, 
fleshy; endocarp thin, hard, bony, breaking up in longitudinal lines during ger- 
mination, tomentose within. 

Dist r . About 180 species in the Neotropics, one species in West Africa; three species in Malesia from the 
Malay Peninsula to New Guinea and the Philippines, but not in the Lesser Sunda Islands. 

Uses. The timber is strong and durable and resistant to marine borers. It is hard to work because of silica. 

Note. The description above is for the Malesian element of Licania; the genus is much more variable in 
the Neotropics. The three Asian species are placed in subgenus Angelesia by Prance & White, Phil. Trans. 
Roy. Soc. London 320 (1988) 94. 


1. Fruit 1-1.3 cm long, ellipsoid, not narrowed towards base or apex. 

2. Leaves oblong, the apices distinctly acuminate; inflorescence rachis densely puberulousl. L. splendens 

2. Leaves elliptic to oblong-elliptic, the apices acute to rounded and emarginate; inflorescence rachis sparsely 

puberulous 2. L. palawanensis 

1 . Fruit 2.5-5 cm long, narrowed at apex and base, fusiform 3. L. fusicarpa 

1. Licania splendens (Korth.) Prance, FI. Neotrop. 
9 (1972) 172. - Angelesia splendens Korth. Ned. 
Kruidk. Arch. 3 (1854) 384; Boerl. & Koord. Ic. 
Bog. 1, 4 (1901) 59, t. 96; Merr. Philip. J. Sc. 10 
(1915) Bot. 307; Enum. Philip. PI. 2 (1923) 236; Cor- 
ner, Wayside Trees (1940) 526; Browne, For. Trees 
Sarawak & Brunei (1955) 307. - Licania angelesia 
Blume, Melang. Bot. 2 (1855) 358. - Chrysobalanus 
splendens Korth. ex Miq. F1. Ind. Bat. 1, 1 (1855) 
358, in syn. - Parinarium fragile Teijsm. & Binn. 
Cat. Hort. Bog. (1866) 253, nom. nud. - Parinarium 
nitidum Hook. /. Fl. Brit. India 2 (1878) 310. - 
Ferolia nitida {Hook, f.) Ridley, J. Str. Br. Roy. As. 
Soc. n. 82 (1920) 183; Fl. Mai. Pen. 1 (1922) 671. - 
Parinarium philippinense Elmer, Leafl. Philip. Bot. 
10(1939)3809. - Fig. 2. 

Tree to 25 m tall, the young branches sparsely Ian- 
ate, soon glabrous. Stipules linear-lanceolate, to 3 
mm long, caducous. Leaves 4-11 by 1.8-4.2 cm, 
oblong, usually acuminate at apex, cuneate at base, 
glabrous beneath; petioles 2-5 mm, canaliculate, 
glabrous when mature. Inflorescence terminal and 
axillary panicles of cymules, 1.5-14 cm long, the 
rachis and branches grey-puberulous. Flowers c. 2 
mm long. Receptacle campanulate, slightly swollen 
to one side, grey-tomentellous on exterior, tomen- 
tose within; pedicels c. 1 mm long. Calyx lobes acute, 
tomentellous on both surfaces. Petals pubescent on 
exterior. Stamens 7-10, slightly unilateral, the fila- 
ments glabrous. Ovary at or near base of receptacle. 

unilocular, pilose on exterior. Fruit ellipsoid, 1-1.3 
cm long; epicarp smooth, glabrous; mesocarp thin, 
fleshy; endocarp thin, hard, bony, breaking open by 
longitudinal lines of weakness, tomentose within. 

Distr. Thailand; in Malesia: Sumatra, Malay 
Peninsula, W. Java, Borneo, Philippines. Fig. 3. 

Ecol. Commonest in forest, including diptero- 
carp forest, on hill slopes and ridges, but wide-rang- 
ing in peat swamp, freshwater swamp forest, on sea- 
shores, and in rocky places; 0-400(-800) m altitude. 

Uses. The timber is strong, durable and resistant 
to marine borers and is used for saltwater piles, rail- 
road ties, etc. However, it is extremely hard to work 
and requires special tools because of silica. The fruit 
is edible but is not widely used. 

Vern. Malay Peninsula: champrai, medang 
merah, m. puteh, membatu, mempadang, merbatu 
kechil; Borneo: piasau-piasau, Kedayan, gandulong, 
Dusun, tampaluan, Sabah, sampaluan, Brunei, 
buku-buku, bunga, djentihan burung, mauhi, 
Kalimantan; Philippines: taguilom bay; amayan, ba- 
lik, D.Bis., dagingan, dagingdingan, S.L.Bis., 
gapas, maralibus, Tagb. 

2. Licania palawanensis Prance, Brittonia 31 (1979) 

Shrub, young branches sparsely puberulous soon 
becoming glabrous. Stipules lanceolate, 1-2 mm 
long, glabrous, caducous. Leaves 3-6 by 1.4-3 cm. 


Chrysobalanaceae (Prance) 


Fig. 3. Distribution of Licania splendens (Korth.) Prance (dots) and L. palawanensis Prance (triangles). 

elliptic to oblong elliptic, rounded to acute at apex, 
emarginate, subcuneate at base, glabrous beneath; 
petioles 1-3 mm long, c. 1.5 mm wide, lanate be- 
coming glabrous with age, rugose. Inflorescences 
panicles of cymules, 3-4 cm long, the rachis and 
branches sparsely puberulous. Flowers c. 2 mm long. 
Receptacle campanulate, slightly swollen to one side, 
grey-tomentellous on exterior, tomentose within; 
pedicels c. 1 mm long. Calyx lobes acute, tomentel- 
lous on exterior, puberulous within. Petals puberu- 
lous on exterior. Stamens 7, inserted around com- 
plete circle, the filaments glabrous. Ovary inserted at 
base of receptacle, lanate-pilose, unilocular. Fruit 
(immature) ellipsoid, epicarp smooth, glabrous; 
mcsocarp thin; endocarp thin, hard, bony, tomen- 
tose within, breaking open by longitudinal lines of 

Distr. Malesia: Philippines (Palawan). Fig. 3. 

Ecol. Confined to ultrabasic rock formation; 
0-3(X) m altitude, including sea-shore forest. 

V I.icania fuMcarpa (KosTkRM.) Prance, Brittonia 
19 (1987) 3W). Hunga fusicarpa Kostfrm. Rcin- 
wardtia 10(1985) 123. 

Tree to 7 m tall, young branches puberulous, gla- 
brescent, with small prominent round lenticels. Stip- 
ules not seen. Leaves 5-10 by 1-4.5 cm, charta- 
ceous, oblong to elliptic, acute to bluntly acuminate 
at apex, cuneate at base, glabrous and glossy on both 
surfaces, decurrent onto petiole; petioles 2-3 mm 
long, rugose, puberulous becoming glabrous with 
age. Inflorescences terminal and axillary panicles of 
cymules, few-flowered, the rachis and branches 
sparsely puberulous. Flowers c. 2 mm long. Recep- 
tacle campanulate, grey-tomentellous on exterior, 
tomentose within; pedicels 2-3 mm long. Calyx 
lobes acute, narrow, tomentellous on exterior, pu- 
berulous within. Petals not seen. Stamens persistent 
beneath young fruit, 0.5-1 mm long, connate at 
base. Fruit (2.5-)3-5 cm long, narrowly spindle- 
shaped, narrowed at apex to a tip 2-3 mm long, nar- 
rowed at base in stipe 5-10 mm long; epicarp 
smooth, glabrous; mesocarp thin; endocarp hard, 
bony, c. i mm thick, densely lanate within, without 
lines of dehiscence. 

Dislr. Malesia: E. Papua New Guinea (Milne 
Bay Prov., [-erguson I., Morobc Prov.). Fig. 5. 

Ecol. Coastal rain-forest, 0-300 m altitude. 


Flora Malesiana 

[ser. I, vol. 10^ 

F\g. 4. Paraslemon urophyllus (V^ ALL. ex A.DC.) A.DC. A. Hab'n, x 0.5; B. flower, x 10; C. fruit, xl;D. 
flower section, x 10 (A, B Sinclair 39504, C, D Sinclair 3319). 


A.DC. Ann. Sci. Nat. Bot. ser. 2, 18 (1842) 208; Miq. F1. Ind. Bat. 1, 1 (1855) 
359; Hook./, in Benth. & Hook./., Gen. PI. 1 (1865) 607; Fl. Brit. India 2 (1878) 
312; BoERL. Handl. Fl. Ned. Ind. 1 (1890) 426; Focke in E. & P. Nat. Pfl. Fam. 
3, 3 (1891) 60; Merr. Philip. J. Sc. 10 (1915) Bot. 307; Ridley, Fl. Mai. Pen. 
1 (1922) 672; Merr. & Perry, J. Arn. Arb. 21 (1940) 197; Corner, Wayside 
Trees (1940) 526; Hutch. Gen. Flow. PI. 1 (1964) 193. - Diemenia Korth. 
Ned. Kruidk. Arch. 3 (1854) 388; Boerl. Handl. Fl. Ned. Ind. 1 (1890) 425. - 
Trichocarya Miq. Fl. Ind. Bat. 1, 1 (1855) 357, p.p. - Fig. 4. 

Tree or shrub. Stipules small and triangular, caducous. Leaves glabrous on 
both surfaces, without stomatal cavities, with 2 small discoid glands at base of 
lamina; petioles eglandular. Inflorescence an axillary or rarely terminal simple 
or sparsely branched raceme. Bracts and bracteoles small, eglandular, not en- 
closing groups of flower buds. Flowers hermaphrodite or polygamo-dioecious. 
Receptacle patelliform or shallowly cupuliform, shortly hairy within; calyx 
lobes 5, acute, subequal. Petals 5, not exceeding calyx lobes, not clawed. Sta- 
mens either 5 and all fertile or 2 fertile with 3 staminodes; the filaments 
glabrous, shorter than the calyx lobes. Ovary centrally inserted at base of recep- 


Chrysobalanaceae (Prance) 


tacle, glabrous or densely hairy on exterior; carpel unilocular, with 2 ovules. 
Style filiform, puberulous towards the base, with 3 large undivided lobes at apex 
or 1 obscure lobe and 2 large, sometimes deeply divided lobes. Fruit a small 
drupe to c. 1.5 cm or c. 3 cm long; epicarp smooth, not lenticellate; endocarp 
thin, hard, bony, smooth on exterior, glabrous within; with 2 large lateral plates 
which break away on germination to allow seedling escape. 

Distr. Three species; Nicobar Islands; in Malesia: Malay Peninsula, Sumatra, Borneo, Moluccas, New 
Guinea, Admiralty Is. 


1. Fruit 1-1.5 mm long. Primary veins of leaves 8-11 pairs. 

2. Receptacle shallowly cup-shaped, glabrous on exterior; fertile stamens 2; style 3-lobed at apex. Inflores- 
cence glabrous 1. P. urophyllus 

2. Receptacle saucer-shaped, puberulous on exterior; fertile stamens 5; style with one obscure and two large 

bifid apical lobes. Inflorescence villous 2. P. versleeghii 

1. Fruit 2.3-3.5 mm long. Primary veins of leaves 5-6 pairs 3. P. grandifnictus 

1. Paraslemon urophyllus (Wall, ex A. DC.) A. DC. 

Ann. Sci. Nat. Bot. ser. 2, 18 (1842) 208; MiQ. Fl. 
Ind. Bat. 1, 1 (1855) 359; Boerl. & Koord. Ic. Bog. 
1, 4 (1901) 61, t. 97; Ridley, Fl. Mai. Pen. 1 (1922) 
672; BuRK. Diet. (1935) 1693; Corner, Wayside 
Trees (1940) 526; Browne, For. Trees Sarawak & 
Brunei (1955) 308; Kochum. & Wyatt-Smith, Mai. 
For. Rec. 17 (1964); Prance & Whitm. Tree Fl. 
.Malaya 2 (1973) 331. - Embelia urophyl/a [Wall. 
Cat. (1830) n. 2309, nom. nud.] ex A. DC. Trans. 
Linn. Soc. 17 (1837) 131. - Diemenia racemosa 
(KoRTH.) MiQ. Fl. Ind. Bat. 1, 1 (1855) 358. - 
Licania diemenia, Melang. Bot. 2 (1855) 10; 
Hassk. Flora 41 (1858) 256, nom. illeg. - 
Paraslemon spicatus Ridley, J. Str. Br. Roy. As. 
Soc. n. 75 (1917) 29. - Fig. 4. 

Tree to 40 m tall, or shrub, the young branches 
glabrous, the trunk often buttressed. Stipules 
triangular, c. 1 mm long, caducous. Leaves thinly co- 
riaceous, narrowly oblong, 2.5-8 by 1.4-2.5 cm, 
cuspidate acuminate at apex, the tip 5-15 mm, 
cuneaic at base; midrib plane above, prominulous 
beneath; primary veins 8- 1 1 pairs; petioles 4-5 mm 
long, canaliculate, glabrous. Inflorescence of ax- 
illary and rarely terminal racemes or occasionally 
slightly branched, 4- 14 cm long, the rachis glabrous. 
Flowers polygamo-dioecious, c. 1 .5 mm long. Recep- 
tacle broadly cupuliform to flattened sauccr-shapcd, 
glabrous on cxicrior, lomcntosc within; pedicels up 
to 2 mm long. Calyx lobes acute, glabrous on exte- 
rior. Petals 5. Stamens 2 fertile and 3 sterile stami- 
nodcs opposite. Ovary inserted at base of receptacle, 
pilose on exterior, unilocular. Style pilose at base, 
glabrous above, the stigma trifid. fruit ellipsoid, 
1-1.5 cm long; epicarp smooth, glabrous; mcsocarp 

thin, hard; endocarp thin, hard, bony, glabrous 
within, opening by 2 lateral plates. 

Distr. Nicobar Islands; in Malesia: Malay Penin- 
sula, Sumatra, Borneo. Fig. 5. 

Ecol. Characteristic of peat swamp forest where 
it is a common large tree, but wide ranging into 
shorter, more open scrub forest. 

Uses. The wood is hard to use because of the 
silica content, but it is used locally for general con- 
struction, posts, and as firewood. 

Vern . Malay Peninsula: kelat, k. pasir, k. puteh, 
nylas; Sumatra: galam tabanga, kayu gelang, malas, 
meriawak; Borneo: mandailas, Brunei, Besaya, sem- 
palawan, Brunei, tempalawan, Bajau, mengilas, 
ngilas padang, obah, Sarawak. 

Notes. The only record of this species from Java 
(Blume s.n., L) is very dubious since the collector's 
name was added later. It is probably either misla- 
belled or from cultivated material. The only dif- 
ference given between P. spicatus and P. urophyllus 
is that the former is a shrub with sessile flowers. 
Some forms of P. urophyllus have extremely short 
pedicels and most sessile-flowered individuals are 
recorded as being small trees. There is thus no reason 
to maintain P. spicatus as a distinct species. 

2. Paraslemon versleeghii Merr. & Pi:rrv, J. Arn. 
Arb. 21 (1940) 197. 

Tree to 40 m tall, the young branches sparsely 
puberulous, soon glabrous. Stipules triangular, c. I 
mm long, caducous. Leaves thinly coriaceous, nar- 
rowly oblong. 5-9.5 by 1.8-3.7 cm, cuspidate acu- 
minate at apex, the tip 7-15 mm long, cuncaie at 
base; midrib plane above, prominulous beneath; 
primary veins 8-12 pairs, inconspicuous, slightly 


Flora Malesiana 

[ser. I, vol. 104 


Fig. 5. Distribution of Licania fusicarpa (Kosterm.) Prance (diamond), Parastemon urophyllus (Wall, ex 
A. DC.) A. DC. (dots), P. grandifructus Prance (squares), and P. versteeghii Merr. & Perry (triangles). 

prominulous beneath; petioles 3-7 mm long, terete, 
glabrescent. Inflorescence of axillary and terminal 
racemes, 2-9 cm long, the rachis sparsely villous. 
Flowers hermaphrodite, c. 1.5 mm long; pedicels 
1.5-3 mm, sparsely villous. Receptacle broadly 
cupuliform-flattened, sparsely villous on exterior, 
tomentose within. Calyx lobes acute, with hirsute 
margins. Petals 5, with hirsute margins. Stamens 5, 
all fertile, opposite the petals in a complete circle. 
Ovary inserted at base of receptacle, glabrous on ex- 
terior except at base, unilocular. Style pilose at base, 
glabrous above, the stigma with two large apical 
lobes, the third reduced or missing. Fruit ellipsoid, c. 
1.6 cm long; epicarp smooth, glabrous; mesocarp 
thin, hard; endocarp thin, hard, bony, glabrous 
within, opening by two lateral plates. 

Distr. Malesia: Moluccas (Morotai), New 
Guinea, and Admiralty Is. Fig. 5. 

E c o 1 . Usually in well-drained hill forest up to 700 m 
altitude, but also reported from secondary forest, 
swampy (Campnosperma) forest and even beach 

Vern. New Guinea: mangu, Tobelo, noeng, 
Irian, gorsauw. Tor, gwarsau, Wainlag, sirebo, 
sisero, Kemtoek, sosopi, Japen, telek, Mooi, sino- 
ree, Biak. 

3. Parastemon grandifructus Prance, Brittonia 39 
(1987) 366. 

Tree to 30 m tall, the young branches glabrous, the 
trunk lightly buttressed to 1 m. Stipules caducous 
(not seen). Leaves coriaceous, narrowly oblong, 
5-8.5 by 1.8-3.2 cm, with long cuspidate acumen at 
apex, the tip 10-16 mm long, cuneate at base, 
glabrous on both surfaces; midrib prominent above, 
prominulous or plane beneath; primary veins 5-6 
pairs, prominulous above, plane beneath; petioles 
5-8 mm long, glabrous, slightly canaliculate, slight- 
ly swollen at base. Inflorescence of axillary and ter- 
minal racemes, the rachis glabrous. Flowers seen on- 
ly in fruiting specimens. Calyx lobes 5, acute, 
glabrous on exterior, glabrous within except for a 
few hairs around base. Receptacle glabrous on ex- 
terior in fruiting condition. Style persistent below 
fruits, the stigma bifid or trifid. Fruit ellipsoid, 
2.3-3.5 by 1.3-1.5 cm, epicarp smooth, glabrous; 
mesocarp thin, 0.25 mm; endocarp thin, hard, bony, 
0.25 mm thick, glabrous within, opening by 2 lateral 
plates 1.9-2 cm long. 

Distr. Malesia: Borneo (Sarawak, Sabah). Fig. 

Ecol. Upland white sand areas. 

Vern. Borneo: ngilas, Ihan, praus, Dyak. 


Pancher e-v Prance, Brittonia 31 (1979) 79; FI. Nouv. Caled. et Dep. 12 (1983) 
106. - Fig. 6, 8. 
Shrubs or small trees. Stipules lanceolate and persistent (absent or very early 

1989] Chrysobalanaceae (Prance) 651 

caducous in New Caledonian species). Leaves usually glabrous on both surfaces 
(lanate beneath in some New Caledonian species), with a pair of, often obscure, 
marginal glands towards the base, without stomatal cavities; petioles eglan- 
dular. Inflorescence a few-flowered terminal or axially raceme of cymules. 
Bracts and bracteoles small, persistent, not enclosing the flowers in small 
groups. Flowers hermaphrodite, slightly zygomorphic. Receptacle campanu- 
late, slightly asymmetric, shortly puberulous on exterior, densely pubescent 
within. Calyx lobes 5, acute. Petals 5, small, not exceeding calyx lobes, not 
clawed. Stamens 5-9, not exceeding calyx lobes, unilateral with 3-7 stami- 
nodes opposite. Ovary inserted midway up receptacle, densely hairy on exterior; 
carpel bilocular with one ovule in each loculus. Style truncate but distinctly 3- 
lobed at apex. Fruit small, fleshy, bilocular or often with one loculus underde- 
veloped; epicarp smooth, not ridged, not lenticellate; mesocarp thin, fleshy; en- 
docarp thin, hard, bony, with a smooth surface, interior very hairy, with 4-6 
longitudinal lines of weakness which allow the seedling to escape. 

Distr . There are 1 1 species, 8 of which occur in New Caledonia and the Loyalty Is., 3 in Malesia: Papua 
New Guinea. 


1. Inflorescence branches glabrescent; flowers glabrescent on exterior. Leaves with conspicuous anastomos- 
ing venation, oblong-elliptic to elliptic, 4-8.5 cm broad. 

2. Leaves elliptic, 7.5-8.5 cm broad 1. H. novoguineensis 

2. Leaves oblong-elliptic, 4-6.5 cm broad 2. H. papuana 

1 . Inflorescence branches lanate to puberulous; flowers pubescent on exterior. Leaf with venation not con- 
spicuously anastomosing, oblong-lanceolate, 2-3.7 cm broad 3. H. longifolia 

1. Hunga novoguineensis Prance, Brittonia 31 2. Hunga papuana (Baker/.) Prance, Brittonia 31 

(1979) 88. - Fig. 6 G, H. (1979) 88. - A ngelesia papuana Baker/. J. Bot. 61, 

Tree 4 m tall, the young branches puberulous, Suppl. (1923) 13. - Fig. 6 A-F. 

soon glabrous, lenticellate. Stipules lanceolate, pu- Small tree, the young branches lanate, soon gla- 

berulous, c. 5 mm long, persistent. Leaves coria- brous. Stipules lanceolate, 3-6 mm long, puberu- 

ceous, elliptic, 15-19 by 7.5-8.5 cm, glabrous on lous, persistent. Leaves coriaceous, oblong-elliptic, 

both surfaces, apex acuminate, the acumen 8-10 10-19 by 4-6.5 cm, finely acuminate at apex, the tip 

mm long, subcuneate at base; primary veins 11-14 8-16 mm, rounded to subcuneate at base, glabrous 

pairs, anastomosing 4 mm away from margins, pro- on both surfaces; primary veins 9-13 pairs, promin- 

minulous above, prominent beneath; petioles 5-6 ulous above, prominent beneath, conspicuously ana- 

mm long, puberulous soon becoming glabrous, stomosing 5 mm from margin; petioles 2-4 mm 

slightly canaliculate, eglandular. Inflorescence of long, shallowly canaliculate, sparsely puberulous 

terminal and axillary panicles, the rachis and bran- when young, soon glabrescent, rugose. Inflores- 

ches puberulous. Bracts and bracteoles 1-2 mm cences of terminal and axillary panicles, 3-10 cm 

long, puberulous, persistent. /^/ow^ri not seen. Frw;7 long, the rachis and branches sparsely lanate- 

sagiltalc pyriform, c. 3.5 cm long, the upper part puberulous when young. Bracts and bracteoles 

triangular, 2-2.5 cm long, the base with a stipe 6- 10 0.5-2 mm long, sparsely puberulous-glabresccnt on 

mm long; epicarp glabrous, smooth, mesocarp thin, both surfaces. I'lowers 1.5-2 mm long. Receptacle 

fleshy; endocarp thin, hard, bony, lanale within. campanulatc, glabrous externally, tomentosc within. 

Pistr. Malesia: Papua New Cjuinca, two collcc- Calyx lobes glabrous on both faces except for cilio- 

tions, from Morobc and Milne Bay Prov. Fig. 7. late margins. Petals glabrous. Stamens c. 7, 

Ecol. Oak forest on slopes, at c. 800 m altitude. unilateral with toothed staminodcs opposite. Ovary 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 6. Hunga papuana (Baker f.) Prance. A. Habit, xO.5; B. flower, x7; C. flower section, x9; D. petal, 

X 15; E. fruit, xO.5; F. ovary section, x 15. - H. novoguineensis Prance. G. Young fruit section, x 1; H. 

habit, xO.5 (A-C Forbes 504, £)-F Womersley NGF 19307, G, H Hartley 12645). 


Chrysobalanaceae (Prance) 




£=» ^ 










1 \ 

Fig. 7 

. Distribution of Hunga longifolia Prance 
H. novoguineensis Prance (dots), and H. pa- 
puana (Baker/.) Prance (triangles). 

bilocular, lanate-pilose externally. Style pilose at 
base, glabrous above, stigma truncate. Fruit sagit- 
tate-pyriform, unilocular, to 5 cm long, the upper 
portion triangular, 2-3.5 cm long, the base narrowly 
and abruptly tapered to a stipe 0.6-1 cm long; 
epicarp smooth, glabrous; mesocarp thin, fleshy; en- 
docarp thin, hard, bony, lanate-tomentose within. 

Distr. Malesia: Papua New Guinea. Fig. 7. 

Ecol. Oak forest; 500-l(X)0 m altitude. 

3. Hunga longifolia Prance, Brittonia 31 (1979) 84. 
- Fig. 8. 

Tree 15 m tall, the young branches puberulous, 
soon glabrous. Stipules linear-lanceolate, 5-6.5 mm 
long, puberulous, subpersistent. Leave.s coriaceous, 
oblong-lanceolate, 7-13 by 2-3.7 cm, acute to 
bluntly acuminate at apex, cuneate at base, glabrous 
on both surfaces; petioles 3-5 mm long, shallowly 
canaliculate, lanate when young, glabrescent with 


vT Zokm 

Fig. 8. Hunga longifolia Prance. A. Habit, xO.5; B. flower; C. petal; D. ovary section, all x20. 

654 Flora Malesiana [ser. I, vol. 10^ 

age, slightly rugose. Inflorescences axillary and ter- forciliate margins. Stamens6-8, unilateral with 3-5 

minal panicles of cymules 1.5-6 cm long, the rachis short staminodes opposite them. Ovary bilocular, in- 

and branches appressed lanate when young, becom- serted midway up receptacle tube, pilose on exterior. 

ing puberulous. Bracts and bracteoles 1-3.5 mm S/^e pilose at base. Fru/7 not seen. 

long, sparsely puberulous-giabrescent on both sur- Distr. Malesia: Papua New Guinea (Misima I.), 

faces. Flowers 2-2.5 mm long. Receptacle cam- known from a single collection. Fig. 7. 

panulate, swollen slightly to one side, lanate-tomen- Ecol . Rain-forest on N. slope, at 300 m altitude. 

tose on exterior, tomentose within. Calyx lobes Fl. July. 

pubescent on both surfaces. Petals glabrous except 


AuBL. Hist. PI. Guiane Fr. 1 (1775) 204; Hauman, Bull. Jard. Bot. Brux. 21 
(1951) 184, quoad subg. Euparinari tantum; Backer & Bakh./. Fl. Java 1 
(1964) 521, p.p.; Hutch. Gen. Flow. PI. 1 (1964) 192, p.p. excl. syn. Maranthes 
etc.; KosTERM. Reinwardtia 7 (1965) 7, excl. syn. Thelira, Ferolia, Mampata et 
Neou; Prance, Fl. Neotrop. 9 (1972) 178; Prance & Whitm. Tree Fl. Malaya 
2 (1973) 332; White, Bull. Jard. Bot. Nat. Belg. 46 (1976) 310; Distr. PI. Afr. 
10 (1976) 327; Fl. Zamb. 4 (1978) 36; Prance, Fl. Venez. 4 (1982) 325; Smith, 
Fl. Vit. Nov. 3 (1985) 44. - Dugortia Scop. Introd. (1777) 217, nom. illeg. - 
Pahnarium Juss. Gen. PI. (1789) 342; Lamk, Encycl. Meth. Bot. 5 (1804) 17; 
St.Hil. Expos. Fam. 2 (1804) 194, p.p.; R.Br, in Tuckey, Nar. Exped. Riv. 
Zaire Cong. (1818) 433; Steud. Nom. (1821) 591; DC. Prod. 2 (1825) 526; Pom. 
Diet. Sci. 37 (1825) 544; Bartl. Ord. Nat. (1830) 406; G.Don, Gen. Syst. 2 
(1832) 478; Meissn. Gen. (1836/42) 102; Benth. Hook. J. Bot. 2 (1840) 211, 
218; Endl. Gen. (1840) 1252, n. 6411; Benth. in Hook., Niger Fl. (1849) 333; 
MiQ. Stirp. Surin. Select. 2 (1850) 7; Blume, Mus. Bot. Lugd.-Bat. 2 (1852) 94; 
Melang. Bot. 2 (1855) 10; Miq. Fl. Ind. Bat. 1, 1 (1855) 352; ibid. (1858) 1084; 
C.MuELL. in Walp., Ann. 4 (1857) 644; Miq. Suppl. Sumatra (1860) 306; 
Benth. Fl. Austr. 2 (1864) 426; Hook. /. in Benth. & Hook./., Gen. PI. 1 
(1865) 607; Miq. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 237; Hook./, in Mart., 
Fl. Bras. 14 (2) (1867) 49; Baill. Hist. PI. 2 (1869) 435, 482; Kurz, For. Fl. Bur- 
ma 1 (1877) 432; Hook./. Fl. Brit. India 2 (1878) 308; Fritsch, Ann. Naturh. 
Mus. Wien 4 (1889) 33; Boerl. Handl. Fl. Ned. Ind. 1 (1890) 421, 424; Focke 
in E. & P. Nat. Pfl. Fam. 3, 3 (1891) 60; King, J. As. Soc. Beng. 66 (1897) 276; 
K. & V. Bijdr. 5 (1900) 332; Bailey, Queensl. Fl. 2 (1900) 524; Brandis, Indian 
Trees (1906) 278; Backer, Schoolfl. Java 1 (1911) 445; Ridley, Fl. Mai. Pen. 

1 (1922) 666; Merr. Enum. Philip. Flow. PI. 2 (1923) 235; Burk. Diet. (1935) 
1693; Corner, Wayside Trees (1940) 527. - Petrocarya Schreb. in Linn. Gen. 
PI. ed. 8, 1 (1789) 245, nom. superfl. - Pahnarium sect. Petrocarya DC. Prod. 

2 (1825) 526; Benth. in Hook., Niger Fl. (1849) 335, p.p. excl. P. glaberrima 
et P. scabra. - Parinarium sect. Neocarya DC. Prod. 2 (1825) 526, p.p. guoadl 
P. excelsum. - Balantium Desv. ^atBuch.-Ham. Prod. PI. Ind. Occ. (1825) 34. 
- Parinarium subg. Petrocarya (DC.) Miq. Fl. Ind. Bat. 1, 1 (1855) 352. -, 

1989] Chrysobalanaceae (Prance) 655 

Lepidocarpa Korth. Ned. Kruidk. Arch. 3 (1854) 385. - Ferolia O.Kuntze, 
Rev. Gen. PI. 1 (1891) 216, p.p. (non Ferolia Aubl.). - Fig. 15. 

Small or large trees or rarely shrubs. Stipules small to large, persistent or ca- 
ducous. Leaves usually with stomatal crypts filled with pubescence on lower sur- 
face or rarely glabrous, or lanate pubescent without crypts. Petioles usually with 
2 circular glands above. Inflorescence a many-flowered complex cyme or cy- 
mose panicle. Bracts and bracteoles eglandular, usually concealing flower buds 
individually and in small groups. Flowers hermaphrodite. Receptacle subcam- 
pulate to cupuliform, slightly swollen to one side, tomentose on both surfaces; 
calyx lobes 5, deltate, acute, densely hairy on both surfaces. Petals 5, as long 
as or shorter than sepals, caducous. Stamens 6-10, unilateral, the filaments 
glabrous, included, with c. 6 minute staminodes opposite. Ovary inserted on up- 
per half of receptacle tube below mouth, pilose on exterior; carpel bilocular with 
1 ovule in each loculus; style arcuate, included. Fruit a fleshy drupe; epicarp ver- 
rucose; endocarp thick, with a rough fibrous surface, with 2 basal obturators 
for seedling escape. 

Disir. Pantropical with 18 species in the Neotropics, 6 in Africa and 15 in tropical Asia (P. anamensis), 
Malesia, the Pacific region (P. insularam) and northern Queensland, Australia; in Malesia 13 species. 

Uses. The fruit of several species are edible, but little-used. 

Note. Since inflorescences and flowers are uniform in the Malesian region, the species are difficult to 
separate; a key containing all 15 Australasian species, based on leaf characters only, is given here. 


1 . Stomatal crypts absent from leaf underside; leaf underside glabrous or with a persistent lanate pubescence 
and then with large persistent stipules 7-40 mm. 
2. Leaf undersurface glabrous. Stipules small and caducous. 
3. Leaves elliptic to oblong or obovate-elliptic, 9.5-20.5 by 4.5-8.5 cm; primary veins 11-16 pairs. Pan- 
icles large and silvery pubescent 1. P. argenleo-sericea 

3. Leaves ovate, 5-9 by 2-4.5 cm; primary veins 7-11 pairs. Panicles small and subsericeous brown 
pubescent 2. P. canarioides 

2. Leaf undersurface densely lanate pubescent, but when removed no stomatal crypts present; stipules large 
and persistent, 7-40 mm long, 3-5 cm broad at base. 

4. Leaves oblong-lanceolate, 5-18 cm long on flowering branches, thickly coriaceous, base cuneate 

3. P. elmeri 

4. Leaves elliptic to oblong, 1 1 -28 cm long, chartaceous, base rounded 4. P. parva 

1. Stomatal crypts present on leaf underside; leaf undersurface lanate or at least pubescent in crypts; stipules 

usually small, or if larger then early caducous. 

5. Leaf lower surface with a series of small glands along lower part of margin. Calyx broadly cupuliform. 
6. Leaf apex acute or rounded but not acuminate; primary veins 10-17 pairs. Young branches with small, 

almost plane icnticels. Low tree of savanna, savanna forest and forest margins 5. P. nonda 

6. Leaf apex acuminate, acumen 3- 10 mm long; primary veins 16-22 pairs; young branches with extreme- 
ly prominent large lenticels; large tree of rain-forest and hills 6. P. papuana 

5. Leaf lower surface without marginal glands on lower part. Calyx usually campanulate. 

7. Primary leaf veins 20-33 pairs (16-26 pairs in P. costata ssp. polyneura). 

8. Petioles 9-17 mm long 7. P. oblonttifolia 

8. Petioles 3-6 mm long. 

9. Leaves chartaceous, primary veins prominulous above, 16-26 pairs 

13c. P. costata ssp. polyneura 

9. Leaves coriaceous, primary veins impressed for upper portion, 20-28 pairs 8. P. gigantea 


Flora Malesiana 

[ser. I, vol. 10^ 

7. Primary leaf veins 20 pairs or fewer. 
10. Petioles 10-20 mm long; leaves with or without a metallic sheen above. 
II. Leaves with metallic sheen above; petioles 14-20 mm long; leaves 4-9 cm broad. Borneo 

9. P. metallica 

11. Leaves without metallic sheen; petioles 10-12 mm long; leaves 6.5-12 cm broad. New Guinea 

10. P. prancei 
10. Petioles 3-10 mm long; leaves without metallic sheen. 

12. Leaves rigidly coriaceous, often broadest well below mid point; midrib and often primary veins 
lightly impressed on upper surface 11. P. rigida 

12. Leaves thinly coriaceous or chartaceous, usually broadest at or above middle (except in P. in- 
sularum); midrib and primary veins usually plane or prominulous. 
13. Leaves ovate, with long thin acumen, tapering from well below mid point; midrib impressed above. 

Plants of Pacific Islands (Fiji, Tonga, Samoa, Wallis Is.) P. insularum 

13. Leaves elliptic to oblong-lanceolate, tapering from middle or above; midrib usually plane or pro- 
minulous. Plants of Sunda shelf. 

14. Inflorescence predominantly axillary. Leaves broadly elliptic 12. P. sumatrana 

14. Inflorescence terminal and subterminal. Leaves elliptic to narrowly oblong. 

15. Leaf apex rounded to acute. Thailand and Indochina P. anamensis 

15. Leaf apex acuminate. Burma, Malay Peninsula, Indonesia and the Philippines 13. P. costata 

1. Parinari argenteo-sericea Kosterm. Reinwardtia 7 
(1965)47, f. 1; 158. 

Trees to 35 m tall; the young branches glabrous, 
prominently lenticellate. SZ/pw/e^ lanceolate, to 8 mm 
long, tomentose on exterior, early caducous. Leaves 
chartaceous, oblong, elliptic to subovate-elliptic, 
9.5-20.5 by 4.5-8.5 cm, glabrous on both surfaces, 
without stomatal cavities beneath, usually 2 glands 
beneath at base near junction with midrib, acute to 
shortly acuminate at apex, the tip 7-10 mm long, 
rounded at base; midrib lightly impressed above ex- 
cept near base, prominent beneath; primary veins 
11-16 pairs, plane above, prominent beneath, erect- 
patent; petioles 5-9 mm long, eglandular, glabrous, 
rugulose. Inflorescence a lax, much branched, ter- 
minal panicle 9-15 cm long, the rachis and branches 
densely grey sericeous-tomentose; bracts and brac- 
teoles ovate, acute, densely tomentellous on exterior, 
glabrous within except near apex, caducous. Recep- 
tacle campanulate, markedly gibbous, densely grey 
tomentellous on exterior, 2-3 mm long; pedicels 1-3 
mm long. Calyx lobes 2-3 mm long, narrowly ovate, 
densely grey tomentose on exterior, tomentellous 
within. Petals spathulate, 2 mm long, caducous. Fer- 
tile stamens 7-8, base forming a conspicuous fused 
ring with opposite tooth-like staminodes. Ovary 
densely pilose. Style pilose, stigma truncate. Fruit 
ovoid, 7-8 by 4.5-5.5 cm, exocarp densely len- 
ticellate; mesocarp thin, fleshy; endocarp extremely 
hard and thick (1 -8 cm thick), woody, granular, and 
very irregularly ridged, with 2 small loculi in centre, 
densely lanate within. 

Distr. Malesia: North Borneo (Sabah). Fig. 9. 

Ecol. Hillside forest, to 100 m altitude; forest 
along rivers. 

Vern. Berangan, Malay. 

Fig. 9. Distribution of Parinari argenteo-sericea 

Kosterm. (stars) and P. canarioides Kosterm. 


2. Parinari canarioides Kosterm. New & Crit. Mai. 
PI. 3 (1955) 25, t. 12 (For. Dept. Bur. of Planning, 
Bogor, Indonesia); Reinwardtia 7 (1965) 159,/. 2. 
Trees to 60 m tall; trunk buttressed to 2.5 m high; 
young branches sparsely puberulous, glabrescent, 
lenticellate. Stipules linear, acute to 5 mm, hirsute, 
early caducous, present on very young leaves only. 
Leaves chartaceous, ovate, 5-9 by 2-4.3 cm, gla- 
brous on both surfaces when mature, without sto- 
matal crypts beneath, acuminate at apex, the tip 
5-12 mm long, rounded to subcordate at base; 
midrib lightly impressed above, prominent beneath, 
sparsely pubescent when young; primary veins 7-11 
pairs, plane to prominulous above, prominent be- 
neath, arcuate; petioles 3-7 mm, glabrous when 
mature, eglandular or with small rather inconspicu- 


Chrysobalanaceae (Prance) 


ous central glands. /n/Zore^cence^ dense-flowered ax- 
illary panicles to 4.5 cm long, the rachis and branches 
tomentose; bracts and bracteoles persistent, ovate, 
puberulous on e.xterior, caducous. Receptacle cam- 
panulate, 3 mm long, tomentose on exterior; pedicels 
1-2 mm long; calyx lobes elliptic, concave, c. 2 mm, 
acute, sparsely puberulous on exterior, densely to- 
mentellous on interior. Petals elliptic, obtuse, 2 mm, 
tapered to base. Fertile stamens 7-8. Fruit ellipsoid, 
3.5-5 by 1 .5-2.5 cm; epicarp densely to sparsely len- 
ticellate; mesocarp fleshy, 1 mm thick; endocarp 5 
mm thick, hard, marbled, densely lanate within. 

Distr. Malesia: Sumatra, Borneo, Sulawesi, 
Philippines (Palawan). Fig. 9. 

Ecol. Forest extending up to 800 m altitude. 

Uses. The timber is much used, but of poor quali- 
ty. Fruit edible, also eaten by pigs. 

3. Parinari elmeri Merr. Univ. Calif. Publ. Bot. 15 
(1929) 92; Kosterm. Reinwardtia 7 (1965) 161, f. 4; 
Prance & Whitm. Tree Fl. Malaya 2 (1973) 335. 

Trees to 32 m, without buttresses; the young bran- 
ches densely tomentellous, glabrescent, obscurely 
lenticellate. Stipules lanceolate, acute, to 18 mm long 
by 3 mm broad at base, lateral, tomentellous, persis- 
tent. Leaves oblong to oblong-lanceolate, 5-18 by 
1 .5-7 cm, chartaceous to thinly coriaceous, glabrous 
above, densely lanate pubescent beneath, without 
stomatal cavities; acuminate at apex, the tip 5-13 
mm long, subcuneate at base; midrib plane or slight- 
ly impressed and pubescent above when young, 
prominent beneath; primary veins 14-21 pairs, 
prominent beneath, curved at margin; secondary 
nerves more or less parallel forming ladder-like 
reticulation; petioles 1.5-6 mm long, tomentellous. 


Fig. 10. Distribution of Pormor/e/fner/ Merr. (stars) 
and P. parva Kosterm. (dots). 

glandular, but glands often obscured. Inflorescences 
of raceme-like reduced terminal and axillary panicles 
or cymules, 1.7-3 cm long, the rachis and branches 
densely brown tomentose; bracts and bracteoles 
large, 2 mm long, ovate, persistent. Receptacle con- 
ical, gibbous, to 3 mm long, brown-lanate on exte- 
rior, pedicels 0.5-2 mm long. Calyx-lobes ovate, 
acute, 2-3 mm long, lanate on exterior. Petals white, 
oblong-ovate, 2-3 mm long, narrowed to base. Fer- 
tile stamens 7-9 with tooth-like staminodes oppo- 
site. Fruit oblong-ellipsoid, 6.7 by 3.7 cm; epicarp 
sparingly lenticellate. 

Distr. Malesia: Malay Peninsula, Borneo (Sara- 
wak, Brunei, Sabah, NE. Kalimantan), Philippines 
(Mindanao). Fig. 10. 

Ecol. Lowland and hill forest to 900 m, including 
areas on ultrabasic rock. 

Uses. The wood is used for supports of Iban long 

Vern. Borneo: resak, Iban. 

4. Parinari parva Kosterm. Reinwardtia 7 (1965) 52, 
f . 5; 162; Prance & Whitm. Tree Fl. Malaya 2 (1973) 


Tree to 15 m tall, bole often fluted at base, without 
buttresses; the young branches densely tomentellous, 
glabrescent, conspicuously lenticellate. Stipules lan- 
ceolate, lateral, 13-37 mm long, up to 5 mm broad 
at base, persistent, conspicuously reticulate and 
densely tomentose on exterior. Leaves chartaceous, 
oblong to elliptic, 11-28 by 5.5-11 cm, glabrous 
above, densely lanate-arachnoid pubescent beneath, 
the pubescence completely obscuring reticulate ner- 
vation, but without stomatal crypts; finely acumi- 
nate at apex, the tip 3-13 mm long, rounded to sub- 
cordate at base; midrib plane and pubescent above, 
prominent beneath; primary veins 15-23 pairs, ar- 
cuate and anastomosing at margin, plane above, 
prominent beneath, pilose; petioles 5-8 mm, densely 
pale brown pilose, with 2-3 extremely prominent 
glands. Inflorescence of short little-branched termi- 
nal and axillary panicles to 5 cm long, sometimes 
borne on young woody branches, the rachis and 
branches densely pale-brown tomentellous; bracts 
and bracteoles ovate, acute, to 5 mm long, persistent. 
Receptacle campanulate, 3 mm long, pale brown 
tomentose on exterior; calyx lobes acute, ovate- 
lanceolate, 1-1.5 mm long, densely tomentose. 
Petals white. Fruit ellipsoid to narrowly ellipsoid, to 
10 cm long by 3 cm broad; epicarp densely len- 
ticellate, ridged when dry; mesocarp thin and fleshy; 
endocarp thick, hard. 

Distr. Malesia: Malay Peninsula (Kelantan, 
Trengganu, Pahang, Johorc), Sumatra, Borneo. Fig. 

Ecol . Mostly on ridge tops and hillsides to 750 m 


Flora Malesiana 

[ser. I, vol. lO^ 

5. Parinari nonda F.v.M. ex Benth. F1. Austr. 2 
(1864)426; BANKSi&SoL.Bot. Cook's Voy. 1 (1900) 
t. 92; Bailey, Queensl. Fl. 2 (1900) 524; Compreh. 
Cat. Queensl. Pi. (1913) 167; Pulle, Nova Guinea, 
Bot. 8, 2 (1910) 367; Kosterm. Reinwardtia 7 (1965) 
170, f. 1 la, excl. syn. P. papuanum et P. salomonen- 
se. - Ferolia nonda (F.v.M. ex Benth.) O. Ktze, 
Rev. Gen. PI. 1 (1891) 216. 

Trees to 15 m tall, without buttresses, the young 
branches sparsely puberulous, soon glabrous, with 
small prominulous ienticels. Stipules lanceolate, 
membraneous, tomentellous, to 5 mm long, very ear- 
ly caducous. Leaves chartaceous to thinly coria- 
ceous, oblong, 4- 1 1 by 1 .8-4.2 cm, glabrous above, 
with stomatal crypts filled with lanate pubescence 
beneath, rounded to acute (or rarely bluntly acumi- 
nate) at apex, subcuneate at base; midrib plane or 
prominulous, sparsely tomentellous when young 
above, prominent beneath; primary veins 10-17 
pairs, curved at margins; secondary nerves reticulate 
slightly flattened, with a series of marginal glands at 
veins on lower portion; petioles 5-10 mm long, 
tomentellous, terete, with 2-4 prominent, conspic- 
uous glands near mid point. Inflorescence of spread- 
ing terminal and subterminal panicles, 5-11 cm 
long, the rachis and branches rather sparsely grey- 
brown tomentellous; bracts and bracteoles large, 
ovate, 2.5-3 mm long, tomentose, caducous. Recep- 
tacle campanulate, 2-3 mm long, tomentose on ex- 
terior; pedicels 0.5-1 mm long. Calyx lobes triangu- 
lar, acute, c. 1 mm long, tomentose on exterior, 
tomentellous within. Petals 5, white, acute. Fertile 
stamens 7-9, with tooth-like staminodes opposite, 
lanate around base. Ovary villous. Style villous 
lanate on lower portion, glabrous above; stigma 
capitate. Fruit ovoid, epicarp sparingly lenticellate. 

D i s t r . Australia (Queensland and Northern Ter- 
ritory) and in Malesia: southern extreme of Papua 
New Guinea and Irian Jaya. Fig. 11. 

Ecol. Savanna, open forest, forest on rocky 
areas in lowlands. 

Vern. Papua New Guinea: warrem. 

6. Parinari papuana C.T.White, J. Am. Arb. 31 
(1950) 86. - Parinari nonda auct. non Benth.: Kos- 
term. Reinwardtia 7 (1965) 170, p.p. 

Large trees to 40 m tall, buttressed or unbut- 
tressed, the young branches puberulous, soon gla- 
brous, with clusters of large prominent Ienticels with 
central slit. Stipules lanceolate, very early caducous. 
Leaves thickly coriaceous to chartaceous, oblong, 
4-18 by 1.5-6.5 cm, glabrous above, with stomatal 
crypts filled with lanate pubescence beneath, acu- 
minate at apex, the tip 3-10 mm long, rounded to 
subcuneate at base; midrib plane or slightly impress- 
ed and sparsely tomentellous when young above, 
prominent beneath; primary veins 16-22 pairs, 

Fig. 11. Distribution of Parinari nonda F.v.M. ex 

curved and anastomosing at margins; secondary 
nerves reticulate, slightly flattened, with a series of 
marginal glands at vein endings on lower portion; 
petioles 2-8 mm long, tomentellous when young, 
terete or slightly canaliculate, with 2 conspicuous or 
sometimes obscure glands. Inflorescence of terminal 
and subterminal panicles, 2-6 cm long, the rachis 
and branches densely tomentose or tomentellous; 
bracts and bracteoles large, ovate, 2-2.5 mm long, 
tomentose on exterior; pedicels 0.2-1.5 mm long. 
Calyx lobes triangular, acute, c. 1 mm long, tomen- 
tose on exterior, tomentellous within. Petals 5, 
white, acute. Fertile stamens 7-8, with tooth-like 
staminodes opposite, lanate around base. Ovary 
villous, style villous on lower portion, glabrous 
above; stigma capitate. Fruit ovoid, 4-6 cm long; 
epicarp sparingly to densely lenticellate; mesocarp 
thin, fleshy; endocarp hard, thick, marbled, lanate 


1. Leaves coriaceous, 4-11 by 1.9-5 cm. Mature 
fruit c. 6 cm long when dry. Montane 

a. ssp. papuana 
1. Leaves chartaceous, 7-18 by 2.5-7 cm. Lowland. 
2. Fruit sparsely lenticellate, small, c. 4 cm long. 

Leaf base subcuneate b. ssp. salomonense 

2. Fruit densely lenticellate, large, c. 6.5 cm long. 
Leaf base usually rounded c. ssp. whitei 

a. ssp. papuana 

Unbuttressed tree. Leaves thickly coriaceous, 
4- 1 1 by 1 .9-5 cm, subcuneate at base. Mature fruit 
c. 6 cm long when dry; densely lenticellate on ex- 

Distr. Malesia: Northern, Central and Eastern 
Papua New Guinea. Fig. 12. 

Ecol. Mountains, 500-2000 m altitude. Fig. 12. 


Chrysobalanaceae (Prance) 


Fig. 12. Distribution of Parinari papuana C.T. White ssp. papuana (dots), ssp. salomonensis (C.T. White) 
Prance (triangles), and ssp. whitei Prance (stars). 

Vern. Korafe, morni, Aiyura, puwirini, Was- 
kuk. Tor, Anona. 

b. ssp. salomonense (C.T.White) Prance, Brittonia 
39 (1987) 369. - Parinari salomonense C.T.White, 
J. Arn. Arb. 31 (1950) 87. 

Buttressed tree. Leaves chartaceous, 7-12 by 3-7 
cm, subcuneate at base. Mature fruit c. 4 cm long, 
sparsely lenticellate on exterior. 

Distr. Solomon Islands. Fig. 12. 

Ecol . Lowland forest, hillsides and ridges to 300 
m altitude. 

Vern. Malmone, Kwara'ae, nakisi, one one, 
sauialu, susui. 

c. ssp. whitei Prance, Brittonia 39 (1987) 369. 
Unbuttressed tree. Leaves chartaceous, 7-18 by 

2.5-6.5 cm, rounded at base. Mature fruit c. 6.5 cm 
long, densely lenticellate on exterior. 

Distr. Malesia: West Irian and Papua New Gui- 
nea along northern coast from extreme west to east. 
Fig. 12. 

Vern. Lowka, Manikiong, ogelel, Mooi. 

7. Parinari oblont>ifolia Hook. /. Fl. Brit. India 2 
(1878) 309; King, J. As. Soc. Beng. 66 (1897) 279; 
RiDLKY. Agr. Bull. Str. & Fed. Mai. St. 1 (1902) 144; 
Fl. Mai. Pen. I (1922) 668; Foxw. Mai. For. Rec. 3 
(1927) 175; Corner, Wayside Trees (1940) 527; 
Kosterm. Rcinwardtia 7 (1965) 165,/. 8; Prance & 
Whitm. Tree Fl. Malaya 2 (1973) 335. - Ferolia 
ohionfii/olia (HfXJK./.) O. Kt/c, Rev. (icn. PI. 1 
(1891) 216. - Partnanum hurneense Mkrr. Univ. 
Calif. Publ. Bot. 15 (1929) 93. 

1 recs to 40 m tall, trunk low thick buttrcs.scd lo 2 
m. the young branches minutely tomentellous, gla- 

brescent, conspicuously prominently lenticellate. 
Stipules ovate to lanceolate, acute, 3-5 mm, pilose 
on exterior, early caducous. Leaves coriaceous, ellip- 
tic to oblong, 14-23 by 4-9 cm, glabrous above, 
with stomatal cavities filled with grey lanate pubes- 
cence beneath, shortly acuminate at apex, the tip 
3-13 mm long, rounded to subcordate at base; 
midrib plane above, glabrous when mature except at 
base, prominent, glabresceiit beneath; primary veins 
23-35 pairs, erect, plane above, flattened and prom- 
inent beneath; secondary veins prominulous and 
parallel ± ladder-like beneath; petioles 9-17 mm 
long, thick, tomentellous, when young, glabrescent, 
eglandular or glandular. Inflorescences of large, 
spreading terminal panicles, 10-21 cm long by 7-12 
cm broad, the rachis and branches yellow-grey to- 
mentellous; bracts and bracteoles ovate, 3 mm long, 
early caducous. Receptacle campanulate, slightly 
gibbous, 3 mm long, densely grey tomentose on ex- 
terior; pedicels 1-3 mm long; calyx lobes ovate, 
acute, 1.5-2 mm long, unequal, grey tomentose. 
Petals white to bluish, lanceolate to spathulate, nar- 
rowed towards base, c. 2 mm long, glabrous. Sta- 
mens 8-10, with tooth-like staminodes opposite. 
Ovary pilose; style glabrous; stigma truncate. Fruit 
ellipsoid, 5-9 by 3-4 cm, epicarp den.sely len- 
ticellate; mesocarp 1.5-2 mm thick; endocarp hard, 
thick, marbled, 7-13 mm thick, fibrous, densely 
lanate within. 

Distr. Malesia: Malay Peninsula (S. Kelantan to 
Johorc), Sumatra, Borneo (Sabah, Kalimantan). 
Fig. 13. 

Ecol . Lowland rain-forest and beside rivers or in 
valleys extending lo 450 m altitude. 

Vern. Malay Peninsula: hedara hulan, kemalau, 
nwntelor, merhalu; dunfiun hukit, Malay; Borneo: 
mankudar, mengkudu, Kalimantan. 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 13. Distribution of Parinari oblongifolia 

Hook./, (dots), P. gigantea Kosterm. (stars), and P. 

metallica Kosterm. (triangles). 

8. Parinari gigantea Kosterm. Reinwardtia 7 (1965) 
182, f. 19. 

Large trees to 40 m tall, trunk fluted at base, the 
young branches densely lanate pubescent, glabres- 
cent, with conspicuous small lenticels. Stipules lan- 
ceolate, acute, to 25 mm long, caducous, membra- 
neous, densely appressed tomentellous on exterior, 
glabrous within. Leaver coriaceous, elliptic, 9-17 by 
5-8 cm, glabrous above, with dense conspicuous 
stomatal crypts beneath, bluntly acuminate at apex, 
the tip 3-6 mm long, rounded at base; midrib plane 
above, prominent and pilose, glabrescent beneath; 
primary veins 20-28 pairs, slightly impressed on up- 
per portion, prominulous on lower portion of upper 
surface, straight, erect, parallel; secondary veins ± 
parallel; petiole thick, 4-7 mm long, tomentellous 
when young, with 2 small round glands on mid point 
above. Flowers not seen. Infructescence axillary, 
3-5 mm long. Fruit irregularly ellipsoid, 6.5 cm 
long, 4 cm broad; epicarp densely lenticellate; meso- 
carp fleshy; endocarp hard, bony, irregularly ribbed, 
lanate within. 

D i s t r . Malesia: Borneo (W. Kalimantan, Sabah). 
Fig. 13. 

Ecol. Lowland forest. 

Vern. Lempong, Kalimantan. 

9. Parinari metallica Kosterm. Reinwardtia 7 (1965) 
49, f. 3; 160, f. 3. 

Trees to 16 m tall, unbuttressed, the young bran- 
ches appressed strigose, glabrescent, conspicuously 
lenticellate. Stipules ovate-lanceolate, acute, 8-15 
mm long, densely brown tomentose, membraneous, 
early caducous. Leaves thickly coriaceous, elliptic, 
8-17 by 4-9 cm, glabrous and shiny with metallic 
sheen above when dry, with dense stomatal crypts 
filled with hairs, apex rounded to shortly blunt acu- 
minate, the tip 0-3 mm long, rounded or subcuneate 

at base; midrib plane above, prominent beneath; 
primary veins 10-15 pairs, prominulous to plane 
above, prominent beneath, erect, curved only at 
margin; petioles 14-20 mm long, glabrescent, with 
inconspicuous glands near to lamina base, puberu- 
lous, glabrescent. Inflorescence of axillary little- 
branched panicles, 4-10 cm long, the rachis and 
branches densely brown tomentellous; bracts and 
bracteoles ovate, early caducous. Receptacle cam- 
panulate, slightly gibbous, 2-3 mm long, ferrugin- 
eous pubescent on exterior; pedicels 0.5 mm long; 
calyx lobes lanceolate, acute, 1 mm long, tomentel- 
lous. Petals lanceolate, glabrous. Stamens c. 8 with 
short tooth-like staminodes opposite. Ovary densely 
pilose. Style glabrous, equalling stamens; stigma 
truncate. Fruit not seen. 

Distr. Known only from Brunei, Sabah, and 
Sarawak. Fig. 13. 

Ecol. Forests on well-drained soil, hillsides, 
50-300 m altitude. 

10. Parinari prancei Kosterm. Reinwardtia 10(1985) 

Trees to 25 m tall, the young branches densely 
brown lanate and pilose, lenticellate. Stipules 
caducous (not seen). Leaves rigidly coriaceous, ellip- 
tic, 9-21 by 6.5-12 cm, glabrous and shiny above 
when mature, lanate when young, with conspicuous 
stomatal crypts filled by lanate pubescence beneath, 
broadly apiculate at apex, rounded to broadly sub- 
cuneate at base; midrib ± plane above, prominent 
beneath; primary veins 14-16 pairs, plane or slightly 
impressed above, prominent beneath, arcuate near 
margins, secondary venation parallel and forming a 
ladder-like reticulum; petioles 10-12 mm long, 
densely ferrugineous lanuginose when young, eglan- 
dular. Inflorescences of axillary little branched small 
panicles or racemes, to 3 cm long, the rachis and 
branches densely appressed tomentellous; bracts and 
bracteoles caducous. Receptacle campanulate-cupu- 
liform, 3-4 mm long, appressed tomentellous on ex- 
terior; pedicels 1.5 cm long; calyx lobes triangular, 
acute, 1.5 mm long, tomentellous. F/-U/7 ellipsoid, c. 
4 by 6 cm diam.; epicarp densely lenticellate; meso- 
carp fleshy, 2 mm thick; endocarp woody, very hard 
and thick, marbled, densely lanate within. 

Distr. Malesia: E. Papua New Guinea (Milne 
Bay Prov., Northern Prov.). Fig. 14. 

Ecol. Lowland rain-forest to 400 m altitude. 

11. Parinari rigida Kosterm. Reinwardtia 7 (1965) 
53, f. 6a, b; 163. - Parinari ashtonii Kosterm. Rein- 
wardtia 7 (1965) 53, f. 7; 164. 

Trees to 30 m tall, unbuttressed, the young branch- 
es tomentellous, glabrescent, inconspicuously len- 
ticellate. S//pu/e5 caducous (not seen). Leaves rigidly 
coriaceous, elliptic to oblong ovate, 7.5-23 by 3-8 


Chrysobalanaceae (Prance) 


Fig. 14. Distribution of Pahnari prancei Kosterm. (diamonds), P. rigida Kosterm. (stars), and P. sumatrana 

(Jack) Benth. (triangles). 

cm, those near to inflorescence much smaller than 
others, broadest below mid point, glabrous and 
shiny above, sometimes slightly bullate, the lower 
surface with stomatal crypts filled with pubescence, 
with 2 glandular areas at junction of midrib and 
petiole below, shortly and broadly acuminate at 
apex, the tip 3-17 mm long, rounded or subcordate 
at base; midrib plane or impressed for upper portion 
above, prominent and appressed pilose beneath 
when young; primary veins 13-20 pairs, slightly im- 
pressed above, prominent beneath, slightly curved at 
margins only; secondary venation flattened or 
rounded, parallel; petioles thick, 3-10 mm long, 
grey-pilose pubescent, rugose, with 2 small glands on 
mid point of upper side. Inflorescences of narrow 
terminal panicles to 13 cm long, the rachis and 
branches tomentose; bracts and bracteoles lanceo- 
late, to 2 mm long, early caducous. Receptacle cam- 
panulate, slightly gibbous, 5 mm long, densely vil- 
lous-tomentose on exterior; pedicels c. 1 mm long; 
calyx lobes elongate triangular, 2-2.5 mm long. 
Petals spathulate. Stamens 6-8. Ovary densely vil- 
lous. Style equalling stamens; stigma capitate. Fruit 
irregularly ellipsoid, 5 cm long, to 4 cm diam., 
tapered towards base almost into a stipe; epicarp 
densely Icnticcllate; mcsocarp thin fleshy; cndocarp 
thick, woody, marbled, lanatc within. 

Distr. Malesia: S. Malay Peninsula, Sumatra, 
Borneo (Sarawak, L. Kalimantan). Fig. 14. 

Ecol. Heath and swamp forests, lowland forest; 
0-1400 m altitude. 

12. Parinari sumatrana (Jack) Benth. in Hook., 

Niger Fl. (1849) 335; Blume, Mus. Bot.Lugd.-Bat. 2 
(1852) 97; MiQ. Fl. Ind. Bat. 1, 1 (1855) 353; ibid. 
(1858) 1084; Suppl. Sumatra (1860) 115; ibid. (1861) 
306; C.Muell. in Walp., Ann. 4 (1857) 644; Flora 41 
(1858) 255; Hook. /. Fl. Brit. India 2 (1878) 309; 
Miers, J. Linn. Soc. Bot. 17 (1879) 336; K. & V. 
Bijdr. 5 (19(X)) 340, p.p. excl. P. costatum auct. non 
Blume; Merr. J. Arn. Arb. 33 (1952) 239; Backer 
& Bakh./. Fl. Java 1 (1964) 522, p.p. excl. P. 
costatum; Kosterm. Reinwardtia 7 (1965) 176. - 
Petrocarya sumatrana Jack, Mai. Misc. 2 (7) (1822) 
67 (repr. Calc. J. Nat. Hist. 4 (1843) 165). - Lepido- 
carpa ovalis (KoRTH.) Blume ex Miq. Fl. Ind. Bat. 1, 
1 (1855) 353. - Ferolia sumatrana (Jack) O. Ktze, 
Rev. Gen. PI. 1 (1891) 216. - Parinarium auct. non 
Blume: Backer, Schoolfl. Java I (1911) 445, p.p. - 
Fig. 15. 

Trees to 30 m tall, without buttresses or small ones 
to 50 cm; the young branches densely tomentellous, 
glabrescent, lenticellate. Stipules oblong, to oblong- 
ovate, 5 - 1 2 mm long by 3 - 5 mm wide at base, mem- 
braneous, early caducous, pilose on exterior. Leaves 
chartaccous to subcoriaccous, elliptic to oblong ellip- 
tic, 7 14( 21) by 3-7.5 cm, obtuse to shortly broad 
acuminate at apex, the tip up to 3 mm long, rounded 
to subcordate at base; glabrous and shiny above, 
with deep-set stomatal crypts beneath obscured by 
dense caducous lanatc pubescence when young; 
midrib plane to slightly impressed above, pilose 
towards base, prominent beneath; primary veins 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 15. Parinari sumatrana (Sack) BEt^TH. A . Habit; B. leaf undersurface with pubescence removed in small 
area to show stomatal cavities; C. flower; D. flower section; E. petal; F. ovary and style; G. ovary section; 

H. young fruit (Kostermans 21859). 


Chrysobalanaceae (Prance) 


9-14 pairs, arcuate, prominulous above, prominent 
beneath; petioles 4-8 mm long, with 2 conspicuous 
glands near middle, lightly canaliculate, glabrescent. 
Inflorescence of short axillary panicles 2-6 cm long, 
the rachis and branches brown tomentose; bracts and 
bracteoles membraneous, tomentellous on exterior, 
puberulous within, caducous. Receptacle conical- 
campanulate, 3 mm long, densely pilose on exterior, 
almost sessile. Petals spathulate, bluish. Calyx lobes 
elongate-triangular, 2 mm long, acute, pilose on 
both surfaces. Fertile stamens 8, unequal. Ovary 
densely pilose. S/v/e glabrous, equalling stamens, the 
stigma truncate. Fruit ellipsoid, 4 by 2.5 cm, epicarp 
densely lenticellate; mesocarp 3-4 mm thick; en- 
docarp marbled in cross section, hard, 5 mm thick, 
densely lanate within. 

Distr. Malesia: Sumatra, W. Java. Fig. 14. 

Vern. Java: kanjere badak. 

Note. This species is distinct from others by the 
predominantly axillary inflorescences. The material 
described as Lepidocarpa ovalis has much larger, 
more pointed leaves than most of the collections, but 
KosTERMANS is correct in placing that name in 
synonymy under P. sumatrana. 

13. Parinari costala (Korth.) Blume, Melang. Bot. 
2 (1855) 10; Miq. FI. Ind. Bat. 1, 1 (1855) 354; ibid. 
(1858) 1084; Suppl. Sumatra (1860) 115; C.Muell. 
in Walp., Ann. 4 (1857) 644; Flora 41 (1858) 255; 
Hook. /. Fl. Brit. India 2 (1878) 309; King, J. As. 
Soc. Beng. 66 (1897) 277; Ridley, Agr. Bull. Str. & 
Fed. Mai. St. 1 (1902) 145; Brandis, Indian Trees 
(1906) 278; Burk. J. Str. Br. Roy. As. Soc. n. 73 

(1916) 200; Merr. J. Str. Br. Roy. As. Soc. n. 76 

(1917) 81; Enum. Born. PI. (1921) 290; Ridley, Fl. 
Mal.Pen. I (1922) 666; Merr. Enum. Philip. Fl. PI. 
2(1923)236; Burk. Diet. (1935) 1667; Heyne, Nutt. 
PI. Ned. Ind. ed. 3 (1950) 697; Kosterm. Rein- 
wardtia 7 (1965) 179, f. 17a, b; Prance & Whitm. 
Tree Fl. Malaya 2(1973) 333. 

For further synonyms, see under the subspecies. 

key to the subspecies 

I. Inflorescence and flowers densely ferrugineous 
villous pubescent. Often at high altitudes 

b. ssp. rubJKinosa 
I . Inflorescence and flowers sparsely to densely grey 

or brown appresscd pubescent. Lowlands. 
2. Primary leaf veins 16-26 pairs; mature leaves 
9- 15.7 cm long, oblong (index 2.3-3.65). Fruit 
cxocarp usually densely vcrrucose 

c. ssp. polyneura 

2. Primary leaf veins 10-16 pairs; mature leaves 

5 10.5 cm long, elliptic (index 1.7 2.7), rarely 

oblong. Fruit cxocarp u.sually sparsely vcrrucose 

a. ssp. coslata 

a. ssp. costata. - Lepidocarpa coslata Korth. Ned. 
Kruidk. Arch. 3 (1855) 387; MiQ. Fl. Ind. Bat. 1, 1 
(1855) 354, in syn., sphalm. Lepidocarya costata. - 
Ferolia costata (Korth.) O. Ktze, Rev. Gen. PI. 1 

Tree to 60 m tall, trunk buttressed up to 2 m, the 
young branches densely appressed tomentellous, 
glabrescent, with small conspicuous lenticels. Stip- 
ules lanceolate, membranaceous, 3-7 mm long, 
pilose on exterior, early caducous. Leaver coriaceous 
or rigidly chartaceous, elliptic, subovate-elliptic to 
oblong (leaf index 1.7-2.7), 5-10.5 by 1.8-4 cm, 
glabrous above when mature but with sparse lanate 
covering when very young, with stomatal cavities 
filled with grey lanate pubescence beneath, acumi- 
nate at apex, the tip 3-5 mm long, round to sub- 
cuneate at base; midrib prominulous above, tomen- 
tellous towards base, prominent beneath; primary 
veins 10-16 pairs, arcuate, prominulous above, 
prominent beneath; secondary veins rounded or only 
slightly flattened; petioles 5-9 mm long, slender, 
tomentellous when young, soon glabrous, usually 
eglandular or with 2 inconspicuous median glands. 
Inflorescences of predominantly axillary or terminal 
few-flowered lax panicles to 8 cm long, the rachis and 
branches appressed grey to brown appressed tomen- 
tellous; bracts and bracteoles lanceolate, c. 2 mm 
long, caducous. Receptacle campanulate, slightly 
gibbous, grey-brown pubescent on exterior, 3-3.5 
mm long; pedicels 0.5-1 mm long; calyx lobes ovate, 
acute, 1.5-2 mm long, grey tomentellous on exte- 
rior. Petals white, spathulate, 1.5-2 mm long, cadu- 
cous, glabrous. Stamens 7-8, with small tooth-like 
staminodes opposite, slightly unequal; style gla- 
brous; stigma capitate. Fruit ellipsoid, to 3.5 by 4.5 
cm; epicarp usually sparsely vcrrucose; mesocarp 2 
mm, fleshy; endocarp hard, marbled, 3-5 mm thick, 
fibrous, densely lanate within. 

Distr. Malesia: Malay Peninsula, Sumatra, 
Borneo, Philippines (Mindanao, Culion, Samar). 
Fig. 16. 

Ecol. Lowland forest, hillsides, ridges; altitude 
up to 300 m. 

Vern. Malay Peninsula: kemalau, mambatu, 
merbatu; Borneo: augok, Piak, bugan, Iban. 

b. ssp. rubii(inosa (Ridley) Prance, Brittonia 39 
(1987) 368. Parinarium helferi Hook./. Fl. Brit. 
India 2 (1878) 311, excl. syn. Parinarium .sumatra- 
num sensu Kurz; Brandis, Indian Trees (1906) 278; 
Kostkrm. Rcinwardtia 7 (1965) 175. - Parinari 
ruhiginosa Ridley, J. Sir. Br. Roy. As. Soc. n. 75 
(1917) 29; Fl. Mai. Pen. I (1922) 668; Foxw. Mai. 
For. Rcc. 3 (1927) 175; Burk. Diet. (1935) 1667; 
KosTiJRM. Rcinwardtia 7 (1965) 168. f. 10; Prance & 
Whitm. Tree II. Malaya 2 (1973) 336. Parinarium 
custatum Ulumi. var. rubiginusum Ridley, J. led. 


Flora Malesiana 

[ser. I, vol. 10"* 

Fig. 16. Distribution of Parinari costata (Kosterm.) Blume (diamonds), P. costcta ssp. rubiginosa (Ridley) 
Prance (triangles), and ssp. polyneura (Miq.) Prance (inverted triangles). 

Mai. St. Mus. 6(1915) 143. - Parinari bicolorMERR. 
Philip. J. Sc. 10 (1915) Bot. 309; Enum. Philip. Fl. 
PI. 2 (1923) 235; Kosterm. Reinwardtia 7 (1965) 172, 
/. 12. 

Leaves 4-11.5 by 1.6-4.3 cm, oblong elliptic to 
oblong lanceolate; primary veins 11-19 pairs; pet- 
ioles 4-8 mm long, thickly tomentose. Inflorescence 
dense to lax, ferrugineous villous pubescent. Fruit 
exocarp sparingly lenticellate. 

Distr. Burma; Malesia: Malay Peninsula, Bor- 
neo (Sabah, Sarawak, Kalimantan), Philippines 
(Mindanao, Bucas Grande I.). Fig. 16. 

Ecol . In lower montane forests of Malay Penin- 
sula and Borneo (750-1500 m) and lowland forests 
of the Phihppines. 

Vern. Merbatu, Malay (= Malesian standard 
timber name for various genera); Borneo: mengku- 
dur, Balikpapan. 

c. ssp. polyneura (Miq.) Prance, Brittonia 39 (1987) 
368. - Parinarium polyneurum Miq. Fl. Ind. Bat., 
Suppl. Sumatra (1860) 115; ibid. (1861) 306; Hook. 
/. Fl. Brit. India 2 (1878) 309; King, J. As. Soc. Beng. 
60 (1897) 278; K. & V. Bijdr. 3 (1901) 340; Kosterm. 
Reinwardtia 7 (1965) 167, f. 9a, b; Prance & Whitm. 
Tree Fl. Malaya 2 (1973) 336. - Ferolia polyneura 
(Miq.) O. Ktze, Rev. Gen. PI. 1 (1891) 216. 

Leaves 9-15.7 by 3.7-6.3 cm, oblong (index 2.3 
-3.65); primary veins 16-26 pairs; petioles 3-7 mm 
long, thick, tomentose. Inflorescence lax, inflores- 
cence and flowers with grey appressed tomentellous 
pubescence. Fruit exocarp usually densely verrucose. 

Distr. Malesia: Malay Peninsula (Kelantan, 
Perak, Pahang, Malacca), Singapore, Sumatra, Bor- 
neo. Fig. 16. 

Ecol. Lowland forest and occasionally in hills 
and seasonal swamps. 

1989] Chrysobalanaceae (Prance) 665 

Excluded species 7520] ex Hook. /. Fl. Brit. India 2 (1878) 311; 

KosTERM. Reinwardlia 7 (1965) 178, /. 16. = 
Parinari wallichiana R.Br, [in Wall., Cat. (1832) Dipterocarpus cornutus Dyer (Dipterocarpaceae). 


Rafin. Sylva Tellur. (1838) 153; Kosterm. Reinwardtia 7 (1969) 421; Prance 
& Whitm. Tree Fl. Malaya 2 (1973) 323; Smith, Fl. Vit. Nova 3 (1985) 47. - 
Atunus RuMPH. Herb. Amb. 1 (1741) 171, t. 66; Lamk, Encycl. Meth. 1 (1783) 
329, non Atunus Rumph. (1743); Panigrahi & Purohit, Taxon 32 (1983) 122. 
- Cyclandrophora Hassk. Flora 25^ Beibl. 1 (1842) 47; Steen. Bull. Jard. Hot. 
Btzg III, 17 (1948) 461; Kosterm. Candollea 20 (1965) 118. - Moquilea sect. 
Cyclandrophora (Hassk.) Endl. Gen. PI. Suppl. 3 (1843) 103. - Parinarium 
subg. Cyclandrophora (Hassk.) Blume, Melang. Bot. 2 (1855) 10; repr. Flora 
N.R. 16 (1858) 255. - Parinarium subg. Macrocarya Miq. Fl. Ind. Bat. 1, 1 
(1855) 354. - Parinarium sect. Cyclandrophora (Hassk.) C.Muell. in Walp., 
Ann. (1857) 644. - Entosiphon Bedd. Madr. J. Lit. Sci. ser. 3, 1 (1864) 44. - 
Parinarium subg. Ill Hook. /. Fl. Brit. India 2 (1878) 308, p.p. - Petrocarya 
auct. non Schreb.: Jack, Mai. Misc. 2 (7) (1822) 68 [repr. Hook. Comp. Bot. 
Mag. 1 (1836)220; Calc. J. Nat. Hist. 4(1843) 164]. - Parinari auct. non Aubl. 
(Parinarium auct. non Juss.): Benth. in Hook., Niger Fl. (1849) 333, p.p.; 
Blume, Mus. Bot. Lugd.-Bat. 2 (1852) 94; Benth. in Benth. & Hook./., Gen. 
PI. 1 (1865) 607; Boerl. Handl. Fl. Ned. Ind. 1 (1890) 431, 424; Focke in E. 
& P. Nat. Pn. Fam. 3, 3 (1891) 60; Koord. Exk. Fl. Java 2 (1912) 338; Ridley, 
Fl. Mai. Pen. 1 (1922) 666. - Fig. 19. 

Small to large trees, ultimate shoots with complicated system of divaricate 
branching. Stipules large, prominently keeled, lateral, persistent or subpersis- 
tent. Leaves almost glabrous on both surfaces, often with minute papillae on 
venation giving beaded appearance, without stomatal crypts, with a pair of 
glands on midrib at or near base of lower surface. Petioles eglandular. In- 
florescence a raceme, or sparsely branched, contracted panicle. Bracts and 
bracteoles persistent, eglandular, not enclosing groups of flower buds. Flowers 
hermaphrodite. Receptacle obconical to cylindrical, as long as or exceeding 
calyx lobes, hollow, hairy inside throughout, throat blocked by retrorse hairs. 
Calyx lobes 5, broadly ovate to lanceolate, tomentellous on both surfaces. 
Petals 5, glabrous, exceeding calyx lobes. Stamens 10-20, posterior, inserted 
unilaterally on margin of disk; filaments free, exserted; staminodes forming a 
barely visible denticulate margin to throat. Ovary inserted at mouth of recep- 
tacle tube, pilose on exterior; carpel bilocular with 1 ovule in each loculus. Fruit 
large; epicarp glabrous, densely verrucose-crustaceous; mesocarp transversely 
fibrous; endocarp hard, thick, shortly and sparsely hairy inside, breaking up ir- 
regularly at germination. Cotyledons large and strongly ruminate. Germination 
cryptocotylar, eophylls alternate. 


Flora Malesiana 

[ser. I, vol. 104 

Distr. About 11 species in Southern India, Thailand, E. to Fiji and Samoa in the Pacific; in Malesia 5 
species in the Malay Peninsula throughout Indonesia, and New Guinea. 

Vern. Merbatu, Malay = Malesian standard timber name for various genera. 


(including species of India and the Pacific) 

1 . Leaf apex rounded; primary veins 6-8 pairs. Fiji A. elliptica (Kosterm.) Kosterm. 

1. Leaf apex acuminate or acute; primary veins usually more than 10 pairs. India. Malesia, or Pacific: Fiji, 

only A. racemosa. 
2. Receptacle tube cylindrical and narrow. 
3. Leaves broadly elliptic, 8- 10 cm broad; rounded at base; apex shortly acuminate, the acumen 2-3 mm 

long 1. A. latifrons 

3. Leaves oblong, 2.5-6 cm broad; subcuneate to rounded at base; apex with long thin acumen 4-22 mm 

4. Receptacle 8-13 mm long. Leaf apex long acuminate; base rounded 2. A. nannodes 

4. Receptacle 5-7 mm long. Leaf apex short acuminate, the acumen 3-10 mm long; base cuneate 

3. A. penangiana 
2. Receptacle tube funnel-shaped to campanulate. 

5. Leaves broadly ovate, thickly coriaceous, cordate at base, 4.5-12 cm long 4. A. cordata 

5. Leaves usually elliptic, chartaceous to thinly coriaceous, usually rounded at base (if cordate then ex- 
ceeding 10 cm in length). 
6. Fertile stamens 12-14. Inflorescence sericeous or sparsely pilose. India. 
1 . Inflorescence sparsely pilose. Leaves elliptic-lanceolate, with 8-10 pairs of primary veins. India 

A. indica (Bedd.) Kosterm. 
7. Inflorescence densely sericeous. Leaves lanceolate, with 12-16 pairs of primary veins. India 

A. travancorica (Bedd.) Kosterm. 
6. Fertile stamens 15-20. Inflorescence tomentellous. Not in India 5. A. racemosa 

1. Atuna latifrons (Kosterm.) Prance & White, 
Phil. Trans. Roy. Soc. Lond. 320 (1987) 132. - 
Parinarium latifolium Hend. Card. Bull. Str. Settl. 
7 (1933) 102, nom. illeg., non latifolium Exell. - 
Parinari latifrons Kosterm. Reinwardtia 7 (1965) 54. 
- Cyclandrophora latifolia (Hend.) Prance in 
Kosterm., Candollea 20 (1965) 121. ~ Atuna latifolia 
(Hend.) Kosterm. Reinwardtia 7 (1969) 421. 

Small tree to 5 m tall, the young branches densely 
lanate-tomentellous becoming glabrous, obscurely 
lenticellate. Stipules lanceolate, to 11 mm long, 
acute, keeled, sparsely appressed pubescent. Leaves 
chartaceous, broadly elliptic, 11-13 by 8-10 cm, 
glabrous and shiny above, slightly bullate, glabrous 
beneath except for sparsely pilose venation, apex 
very shortly abrupt acuminate, the acumen 2-3 mm 
long, rounded at base with base contracted into 
petiole; midrib prominent on both surfaces, slightly 
pilose towards base above, pilose beneath; primary 
veins 12-14 pairs, prominulous inset in a groove 
above, prominent and pilose beneath, venation 
prominulous; petioles thick, 5-7 mm long, terete, 
densely brown lanate when young. Inflorescences of 
axillary little-branched panicles or spikes, to 5 cm 
long, densely brown sericeous; bracts and bracteoles 
to 15 mm long, ovate-lanceolate, acute, densely seri- 
ceous on exterior, appressed puberulous within. Re- 
ceptacle tube narrowly cylindrical, 7-11 mm long, 

sericeous on exterior, sessile; calyx lobes lanceolate 
to oblong-ovate, 5-10 mm long, unequal, densely 
sericeous on exterior, tomentellous within. Petals 
obovate narrowed to base, 10-11 mm long. Stamens 
c. 20, inserted on faucal annulus 2 mm high with 
tooth-like staminodes opposite, the filaments 10-12 
mm long. Ovary densely strigose. Style slender, gla- 
brous; stigma truncate. Fruit unknown. 

Distr. Known only from Malay Peninsula on 
Kedah-Perak border. Fig. 17. 

Fig. 17. Distribution of Atuna latifrons (Kosterm.) 

Prance & White (star) and A. cordata Cockburn 

ex Prance (triangles). 


Chrysobalanaceae (Prance) 


2. Atuna nannodes (Kosterm.) Kosterm. Rein- 
wardtia 7 (1969) 422; Prance & Whitm. Tree Fl. 
Malaya 2 (1973) 325. - Parinari nannodes Kosterm. 
Reinwardtia 7 (1965) 50, f. 4. - Cyclandrophora 
nannodes (Kosterm.) Kosterm. & Prance, Can- 
dollea 20 (1965) 122. 

Trees to 20 m, usually smaller, unbuttressed; the 
young branches sparsely appressed hirsutulous- 
strigose, soon glabrous, obscurely lenticellate. Stip- 
ules narrowly lanceolate, acute, 6-12 mm long, 
strigose to glabrous, subpersistent. Leaves thinly co- 
riaceous, oblong-lanceolate, 6.7-19 by 2.5-5.5 cm, 
glabrous on both surfaces, sometimes slightly bullate 
above, long slender acuminate at apex, the acumen 
7-22 mm long, rounded at base; midrib promin- 
ulous above, prominent beneath; primary veins 
10-12 pairs, arcuate, prominulous on both surfaces 
or sometimes prominent beneath; petioles 2-4 mm 
long, glabrescent, eglandular, the lower part swollen, 
usually curved. Inflorescences axillary racemes 3-7 
cm long, the rachis densely sericeous-tomentellous; 
bracts and bracteoles lanceolate, 3-7(-13)mm long, 
persistent, sericeous. Receptacle cylindrical, 8-13 
mm long, densely sericeous on exterior, sessile; calyx 
lobes to 6 mm long, unequal, acute, sericeous on ex- 
terior. Petals white, spathulate to ovate, 8-12 mm 
long, narrowed to base. Stamens 18-20, black to 
purple, the filaments 10-15 mm long, slightly unilat- 
eral with tooth-like staminodes opposite. Style to 15 
mm long, glabrous; stigma capitate. Ovary pilose. 
Fruit ellipsoid, 3-4 by 1.5 cm, slightly tapered to 
base, crustaceous verrucose on exterior; mesocarp 
2-2.5 mm, fibrous, hard, endocarp thin. 

D i s t r . Malesia: Malay Peninsula (Trengganu and 
Pahang southward), Borneo (Sabah, Sarawak). Fig. 

Ecol. Well drained forests to 500 m altitude. 

Vern. Merbatu, Malay. 

Fig. 18. Distribution of>4/i/na/ion/io^e5 (Kosterm.) 

Kostfrm. (iriangles), A. penanffiana (Kosterm.) 
KosTtRM. (dots). 

3. Atuna penangiana (Kosterm.) Kosterm. Rein- 
wardtia 7 (1969) 422; Prance & Whitm. Tree Fl. 
Malaya 2 (1973) 326. - Cyclandrophora penangiana 
Kosterm. & Prance, Candollea 20 (1965) 124. - 
Parinari asperula auct. non MiQ.: King, J. As. Soc. 
Beng. 66 (1897) 2U, p.p. 

Trees to 20 m tall, unbuttressed, the young branch- 
lets glabrescent, obscurely lenticellate. Stipules lan- 
ceolate, acute, to 7 mm long, glabrous, stiff, subper- 
sistent. Leaves thinly subcoriaceous, oblong to 
oblong lanceolate, 3.7-13 by 2-5.5 cm, glabrous on 
both surfaces, acuminate at apex, the acumen 3-10 
mm, cuneate at base; midrib flattened prominulous 
above, prominent beneath; primary veins 10-13 
pairs, arcuate, prominulous on both surfaces; pet- 
ioles 3-5 mm long, eglandular, glabrescent, smooth, 
not swollen or curved. Inflorescences axillary ra- 
cemes 3-7 cm long, the rachis densely appressed 
pilose; bracts and bracteoles sericeous to 10 mm 
long, persistent. Receptacle cylindrical, 5-7 mm 
long, sericeous pubescent on exterior; calyx lobes 
acute, 4-5 mm long, slightly unequal. Stamens c. 20, 
the filaments to 8 mm long with tooth-like stami- 
nodes opposite. Style to 10 mm long, stigma capitate. 
Ovary pilose. Fruit (immature) ellipsoid, epicarp 
crustaceous, verrucose. 

Distr. Malesia: Malay Peninsula (Penang, 
Perak, Johore, Kelantan and Trengganu). Fig. 18. 

Ecol. Well drained forests to 500 m altitude. 

Vern. Membatu, Malay. 

Note. The two species Atuna nannodes and A. 
penangiana are hard to separate. The larger flowers 
of A. nannodes seem consistent and the species gen- 
erally has leaves with a much longer apex. These may 
be one variable species. 

4. Atuna cordata Cockburn ex Prance, Brittonia 
39 (1987) 364. - Atuna cordata Cockbvrh, Trees of 
Sabah 2 (1980) 82, nom. inval. 

Tree to 40 m tall, the trunk often with thick but- 
tresses; young branches glabrescent, inconspicuously 
lenticellate. Stipules to 1 .7 cm long, very early cadu- 
cous. Leaves coriaceous, broadly ovate, 4.5-12 cm 
long, 3-9.5 cm wide, abruptly acuminate at apex, 
the acumen 1-3 mm long, cordate at base, glabrous 
and shiny above, glabrous beneath; midrib promin- 
ulous above, prominent beneath; primary veins 9- 12 
pairs, lightly prominulous above, prominulous and 
glabrous beneath; petioles 1-3 mm long, short and 
thick, glabrous. Inflorescences of terminal and 
subterminal racemes 4-8 cm long, borne in single or 
more often in paired branches, densely tomcntellous 
on exterior, pubcrulous within; bracts and bracteoles 
ovate, tomcntellous, early caducous. Receptacle 5-7 
mm long, conical to campanulatc, tomcntellous on 
exterior, sessile; calyx lobes slightly unequal, tomcn- 
tellous on both surfaces. Petals c. 7 mm long, 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 19. Aluna racemosa Rafin. ssp. excelsa (Jack) Prance. A. Habit; B. leaf undersurface; C. flower bud; 
D. flower section; E. ovary section; F. base of stamens; G. petal; H. fruit {A-G Whitmore 3542, // Agama 



Chrysobalanaceae (Prance) 


)bovate, glabrous. Stamens c. 10, inserted on one 
ide of ring, the filaments 10-12 mm long. Ovary 
lensely pilose. Style slender, hirsutulous on lower 
)ortion. Fruit 6 cm long, 5 cm wide, ovoid; epicarp 
Tustaceous verrucose, mesocarp 5 mm thick, fi- 
)rous, hard, endocarp thin. 

Distr. Malesia: Borneo (Sabah). Fig. 17. 

Ecol. Hill forests on ultrabasic rock. 

>. .\tuna racemosa Rafin. Sylva Tellur. (1838) 153; 
^ERR. Index Rafin. (1949) 136; Kosterm. Reinward- 
ia7 (1969)422. - Fig. 19. 
For further synonyms, see under the subspecies. 


1. Leaves 10-25(-35) cm long, usually elliptic, 
oblong or lanceolate but sometimes ovate, char- 
taceous or thickly coriaceous, the apex long finely 
acuminate, 6-25 mm long; petioles thick. Flowers 
10-17 mm long. Medium to large trees often with 
fluted bole a. ssp. racemosa 

I. Leaves 4.5-12 cm long, usually ovate or oblong- 
ovate, subcoriaceous or coriaceous, the apex 
bluntly acuminate, 3-10 mm long; petioles thin. 
Flowers 8-11 mm long. Large trees with cylin- 
drical bole b. ssp. excelsa 

I. ssp. racemosa. - Atunus alba Rumph. Herb. 
\mb. 1 (1741) 171, t. 66, nan Atunus litorea Rumph. 
4erb. Amb. 3 (1743) 96, t. 63. - Cydandrophora 
\laberrima Hassk. Flora 25 (2), Beibl. 1 (1842) 47; 
bid. 11 (1844) 583; Cat. Hort. Bog. (1844) 269. - 
°arinari glaberrimum (Hassk.) Hassk. Tijd. Nat. 
3es. Phys. 10(1843) 147; C.Muell. in Walp., Rep. 
i (1845/46) 647; in Walp., Ann. 4 (1857) 645; Blume, 
Vlus. Bot. Lugd.-Bat. 2 (1852) 98; Miq. F1. Ind. Bat. 
I. 1 (1855) 355; K. & V. Bijdr. (1900) 338, incl. var. 
anceolatum (Teijsm. & Binn.) K. & V., p.p. quoad 
;pec. Java; Burk. Diet. (1935) 1696; Backer & 
3akh./. FI. Java 1 (1964) 522. - Parinarium sca- 
irum Hassk. Tijd. Nat. Ges. Phys. 10 (1843) 147, 
lomen; Cat. Hort. Bog. (1844) 269, nomen; Flora 27 
1844) 585; C.MuELL. in Walp., Rep. 5 (1845/46) 
>47; in Walp., Ann. 4(1857)645; Blume. Mus. Bot. 
Lugd.-Bat. 2 (1852) 95, p.p.; Mio. Fl. Ind. Bat. 1, 1 
;i855) 354, t. 5; K. & V. Bijdr. 5 (1900) 337. p.p.; 
Backer, School fl. Java (191 1)445; Ridley, Fl. Mai. 
Pen. 1 (1922)669. ~ Parinarium lanceolalumJtusM. 
k Bins. Cat. Hor(. Bog. (1854) 253. 255. nomen. - 
Parmarium amhomense Teijsm. & Binn. I.e. 254, 
nomen. - Parmarium margarata A.Gray. Bot. 
kVilkcs U.S. Expl. Expcd. 1 (1854) 489. I. 55; 
C.Muell. in Walp., Ann. 4 (1857) 646. - Parina- 
num laurinum A.Gray. Bot. Wilkes U.S. Expl. Ex- 
Xd. 1 (1854) 490. I. 55; C.Muell. in Walp.. Ann. 4 
[1857)646; Merr. Philip. J. Sc. 10(1915) Hot. 210; 

Kanehira. Bot. Mag. Tokyo 45 (1931)282. - Petro- 
carya glaberrima (Hassk.) Miers. J. Linn. Soc. Bot. 
17 (1879) 336. - Ferolia glaberrima (Hassk.) O. 
Ktze. Rev. Gen. PI. 1 (1891) 216. - Ferolia scabra 
(Hassk.) O. Ktze. I.e. 216. - Petrocarya scabra 
(Hassk.) Miers, J. Linn. Soc. Bot. 17 (1897) 336. - 
Parinarium elatum King, J. As. Soc. Beng. 66 (1897) 
280; Ridley, Fl. Mai. Pen. 1 (1922) 669. - Parina- 
rium hahlii Warb. Tropenpfl. 6 (1902) 370. - Pari- 
narium mindanaense Perk. Fragm. Fl. Philip. 
(1904) 119. - Parinarium curranii Merr. Philip. J. 
Sc. 4 (1909) Bot. 264. - Parinarium warburgii Perk. 
ex Merr. J. Str. Br. Roy. As. Soc. n. 76 (1917) 82. 
- Cydandrophora elata (King) Kosterm. Candollea 
20 (1965) 122. - Cydandrophora scabra (Hassk.) 
Kosterm. I.e. 126. - Cydandrophora laurina 
(Gray) Kosterm. I.e. 135. - A. elata (King) 
Kosterm. Reinwardtia 7 (1969) 421; Prance & 
Whitm. Tree Fl. Malaya 2 (1973) 324. - Atuna 
scabra (Hassk.) Kosterm. Reinwardtia 7 (1969) 422. 

Trees to 45 m tall, usually smaller, the bole often 
fluted, young branches glabrous or appressed strig- 
ose. Stipules lanceolate, stiff, to 20 mm long, acute, 
glabrous to strigose. subpersistent. Leaves usually 
chartaceous. more rarely stiffly coriaceous, broadly 
ovate, elliptic, oblong or even lanceolate, 10-25 
(-35) by 3.5- 1 1 cm, acuminate at apex, the acumen 
6-25 mm long, rounded to subcordate at base, 
glabrous on both surfaces when mature, sometimes 
sparsely strigose beneath on lower portion when 
young; midrib prominent on both surfaces; primary 
veins 10-13 pairs, prominulous above, prominent 
beneath, straight or arcuate; the venation conspicu- 
ously papillose and often giving leaf a scabrous ap- 
pearance; petioles thick, 3-7 mm long, glabrous or 
pilose glabrescent . Inflorescences of axillary racemes 
or little branched with up to 3 racemose branches on 
short main peduncle, 5-15 cm long, the rachis 
tomentellous to sericeous; bracts and bracteoles 
ovate, acute, to 8 mm long, caducous. Receptacle 
turbinate-campanulate, 5-10 mm long, tomentose 
to sericeous on exterior; pedicels 0.5-1 mm long, 
calyx lobes 4-7 mm long, ovate to ovate-oblong, 
densely tomentellous on both surfaces. Petals ovate- 
oblong, to 10 mm long, blue or white. Stamens 
15-20, pale blue, to 15 mm long with tooth-like 
staminodes opposite. Ovary densely villous. Style 
equalling filaments, stigma small. Fruit ellipsoid to 
subglobose. to 7.5 cm diam.; epicarp crustaceous 
verrucose; mesocarp to 1 1 mm thick, endocarp thin, 
1-3 mm, dcn.scly pilose within. 

Distr. A wide range from Thailand to the 
Pacific: Admiralty, Caroline, and Solomon Islands, 
Fiji. Tonga. Samoa; in Malesia: Malay Peninsula 
(Pcrak). Singapore. Sumatra, Borneo (Sarawak, 
Brunei), Sulawesi. Philippines. Ambon. Ternate. 
Ccram. New Ciuinca, New Hrilain. lig. 20. 


Flora Malesiana 

[ser. I, vol. 10 

Fig. 20. Distribution of A tuna racemosa Rafin. ssp. racemosa (dots) and ssp. excelsa (Jack) Pranc 
(triangles). Atuna racemosa ssp. racemosa also occurs in Tonga, Fiji, and Samoa outside the area shown i 

the map. 

E c o 1 . Usually occurring in well-drained lowland 
or hill forest, up to 600 m altitude, but also found on 
riverbanks, freshwater or brackish swamps and even 
in mangrove. 

Uses. The fruit (cotyledon) is grated and made 
into a putty for caulking canoes, widely used in 
Pacific islands. An oil is extracted from the seeds 
used variously in different areas, e.g. to scent 
coconut oil and for hairdressing. The leaves are used 
to thatch the outside walls of houses in Fiji. The 
wood is used locally for posts and poles, but is not of 
good quality. 

Vern . Jangong, membatu, Malay; kisokka, Jav.; 
Borneo: belibu, senumpol, Iban, kukut, Sarawak, 
merampangi, tatambu, Sabah, torog, Orang Sungei, 
K'tangan; Sulawesi: lomo, Makassar; Philippines: 
aluma, Ceb., botabon, butabul, getabon, Tagb., 
bolga, Bik., pantog-usa, Kuy., pinae, tabontaba, 
takoutaban. Bis., tabong. Bag., tabon-tabon, C. 
Bis., Bik., Mbo., samake, Bug.; New Guinea: 
asikua, asista, Saki, bata-bata, koewao, Kwerba, 
dela, Mooi, kan, Oriomo, low tukwa, lowtukwa, 
Manikiong, mangosowai, Japen; New Britain: latita, 
tita; New Georgia: y//", tavai, tita, Uso; Caroline Is.: 
agaratim, ais, eis, eritem, grihing, Palau, adidi. Yap, 
Solomon Is.: do-omu, oso, saia, tij, Kwara'ae; Fiji: 
makita; Tonga: hea, seea; Samoa: ifi-ifi. 

Note. KosTERMANS included Cyclandrophoi 
glaberrima as a synonym of Atuna excelsa rath( 
than where it is placed here. There seems little doul 
based on the original description and herbariui 
material at Leiden bearing Hasskarl's writing thi 
C. glaberrima is equal to Atuna racemosa ssp. rac 
mosa as defined here. The original description of < 
glaberrima indicates leaves that are far too large f( 
ssp. excelsa. Atuna excelsa was distinguished t 
KosTERMANS by its coriaceous leaves and short pe 
ioles. However, many sheets which he determined { 
A. racemosa have equally short petioles and there 
much variation in leaf texture. Therefore it is n< 
possible to maintain A. elata. Similarly the distin( 
tion of A. scabra was the scabrous texture of tl 
leaves and their more lanceolate shape. Many colIe( 
tions of /4. racemosa are equally scabrous {e.g. LA 
52392 from New Guinea) and there is so much varii 
tion in leaf shape that it would be quite impossible t 
separate /I. scabra on that feature. This was alread 
placed under Parinari glaberrimum by Backer an 
Bakhuizen van den Brink (I.e. 1964). 

b. ssp. excelsa (Jack) Prance, stat. nov. - Petroci 
rya excelsa Jack, Mai. Misc. 2 (7) (1822) 68 [rep: 
Hook. Comp. Bot. Mag. 1 (1836) 220; Calc. J. Na 
Hist. 4 (1843) 164]; Walp. Rep. 2 (1843) 7. - Parint 


Chrysobalanaceae (Prance) 


rium jackianum Benth. in Hook., Niger Fl. (1849) 
335; MiQ. Fl. Ind. Bat. 1, 1 (1855) 356; C.Muell. in 
Walp., Ann. 4 (1857) 644; Hook./. Fl. Brit. India 2 
(1878) 312. - Parinarium aspentlum MiQ. Fl. Ind. 
Bat., Suppl. Sumatra (1860) \\5,nomen; ibid. (1861) 
307, descr.; Hook. /. Fl. Brit. India 2 (1878) 310; 
King, J. As. Soc. Beng. 66 (1897) 281; K. & V. Bijdr. 
(1900) 337, p./?.; Ridley, Fl. Mai. Pen. 1 (1922)670. 

- Ferolia asperula ^iQ.) O. Ktze, Rev. Gen. PI. 1 
(1891) 216. - Ferolia jackiana /Benth.) O. Ktze, I.e. 

- Parinarium spicatum King, J. As. Soc. Beng. 66 
(1897) 279; Ridley, Fl. Mai. Pen. 1 (1922) 669. - 
Parinarium maingayi King, J. As. Soc. Beng. 66 
(1897) 280; Ridley, Fl. Mai. Pen. 1 (1922) 669. - 
Parinarium villamilii Merr. Philip. J. Sc. 10 (1915) 
Bot. 308; Enum. Philip. Fl. Pi. 2 (1923) 236. - 
Cyclandrophora villamilii ^err.) Prance ex 
Kosterm. Candollea 20 (1965) 126. - Cyclan- 
drophora excelsa (Jack) Kosterm. I.e. 128. - 
Cyelandrophora asperula (Miq.) Prance ex Kos- 
term. I.e. 130. - Atuna villamilii (Merr.) Kosterm. 
Reinwardtia 7 (1969) 422. - Aluna exeelsa (Jack) 
Kosterm. I.e. All; Prance & Whitm. Tree Fl. 
Malaya 2 (1973) 324. - Fig. 19. 

Tree to 45 m tall, the trunk buttressed up to 2 m, 
not fluted, the young branches sparsely strigose, 
glabrescent, obscurely lenticellate. Stipules lanceo- 
late, 8-15 mm long, acute, sparsely strigose, subper- 
sistent. Leaves rigidly chartaceous to coriaceous, 
ovate to oblong-ovate or less frequently oblong, 
4.5-12 by 2-5 cm, acuminate at apex, the acumen 
3-10 mm long, subcordate, rounded or subcuneate 
at base, glabrous on both surfaces; midrib prominent 
on both surfaces; primary veins 9-13 pairs, arcuate, 
prominulous above, prominent beneath, the vena- 
tion papillose giving a beaded appearance; petioles 
slender, 3-6 mm long, puberulous, glabrescent or 
glabrous. Infloreseenees of axillary racemes to 7.5 
cm long, or little branched with 2 or more racemose 
branches on short main peduncle, the rachis and 
branches densely short sericeous; bracts and brac- 

teoles oblong, e. 3 mm long, persistent. Reeeptacle 
turbinate-campanulate, 4-7 long, sericeous on ex- 
terior; calyx lobes ovate, equal, to 4 mm, sericeous 
on exterior, tomentellous within. Petals white to 
bluish white, oblong, to 5 mm long, caducous. Sta- 
mens 13-18, to 8 mm long with tooth-like stamin- 
odes opposite. Ovary pilose. Style glabrous, equal- 
ling filaments, glabrous above, stigma small. Fruit 
subglobose to slightly pyriform, 5-7 cm diam. or 
5-7 by 3.5-4.5 cm; epicarp crustaceous, verrucose; 
mesocarp fibrous, 5-8 mm thick, endocarp thin, 
densely pilose within. 

Distr. Malesia: Malay Peninsula (Kedah and 
Trengganu southward), Sumatra, Java, Borneo, N. 
Sulawesi. Fig. 20. 

Ecol. Lowland forests on well drained soils ex- 
tending to 750 m altitude on ridges and hillsides. 

Vern. Malay Peninsula: kemalau utat, merbatu; 
Sumatra: kemiling utan, klappa soepai, pelee kamb- 
ing, salak; Borneo: membatu, Sabah, mahadiu, Ban- 
djar, temalang. 

Notes . Kostermans is probably correct in inter- 
preting Petrocarya exeelsa Jack as the species 
described here. The original description is quite 
detailed and fits this taxon better than any other 

Kostermans treated these two subspecies as 
separate species. They were differentiated by small 
characteristics of leaf shape, the acumen and the 
base. While there do seem to be two elements involv- 
ed in this complex, there is a complete graduation of 
any single character such as leaf length, apex length, 
petiole thickness, leaf shape or fiower size. Ssp. ex- 
eelsa is much commoner in Sundaland and ssp. 
raeemosa in the Sahul shelf and Pacific islands, but 
the two subspecies have considerable geographical 
overlap with ssp. raeemosa occurring sporadically on 
the Malay Peninsula. Since all characters merge and 
are only weakly correlated, these two species are 
reduced to subspecies, a rank more in accord with 
their variational and geographical patterns. 


Blume, Bijdr. (1825) 89; Kosterm. Candollea 20 (1965) 196; Prance, Bol. Soc. 
Brot. s^r. 2, 40 (1966) 183; Brittonia 20 (1968) 203; Fl. Neotrop. 9 (1972) 201; 
Prance & Whitm. Tree Fl. Malaya 2 (1973) 329; White, Bull. Jard. Bot. Nat. 
Belg. 46 (1976) 294; Distr. PI. Afr. 10 (1976) 313; Fl. Zamb. 4 (1978) 41; 
Letouzey & White, Fl. Cameroun 20, Fl. Gab. 24 (1978) 29. - Exitelia Blume, 
Fl. Jav. 1, Praef. (1828) vii, nom. illeg. - Grymania Presl, Epim. Bot. (1851) 
193, p.p. quoad G. salicifolia tantum. - Parinari sect. Sarcostegia Benth. in 
Hook., Niger Fl. (1849) 335, excl. P. jackiana (Petrocarya excelsa). - Parinari 
Mihg. Sarcostegia (Besth.) MiQ. Fl. Ind. Bat. 1, 1 (1855) 355, ^jrcV. P. Jackiana; 


Flora Malesiana 

[ser. I, vol. lO" 

Fig. 21. Maranthes corymbosa Blume. A. Habit; B. leaf base and glands; C. flower and bud; D. flower se( 
tion; E. petal; F. anthers; G. ovary section; H. fruit (A-G Sulit 19, H Sinclair 10687). 

1989] Chrysobalanaceae (Prance) 673 

Hauman, Bull. Jard. Bot. Brux. 21 (1951) 185. - Pahnari subg. Exitelia Bluue, 
Melang. Bot. 2 (1855) 10; Hassk. Flora 16 (1858) 255. - Pahnari sect. Exitelia 
(Blume) C.Muell. in Walp., Ann. 4 (1857) 645. - Fig. 21. 

Medium-sized to large trees. Stipules deltate, intrapetiolar, stiff, caducous. 
Leaves glabrous on both surfaces when mature (or lanate in African species), 
with dense caducous cobweb-like indumentum when young, without stomatal 
crypts; with paired glands at junction of lamina and petiole. Petioles eglandular. 
Inflorescence a many-flowered corymbose panicle. Bracts and bracteoles eglan- 
dular, caducous, not enclosing flower buds in small groups. Flowers herma- 
phrodite. Receptacle 6bcomcdi\, narrowed into pedicel, solid, almost completely 
filled with nectariferous tissue, short tomentose to glabrous on exterior, gla- 
brous within, calyx lobes suborbicular, deeply concave, unequal. Petals 5, not 
clawed. Stamens 25-40, inserted on margin of disk, unilateral with tooth-like 
staminodes opposite to almost in a complete circle; filaments far exserted 
beyond calyx lobes, in a tangled mass. Ovary inserted laterally at mouth of 
receptacle; carpel bilocular with 1 ovule in each loculus. Style pubescent at base 
only, curved upwards, exserted. Fruit a large fleshy drupe; epicarp smooth, 
glabrous, not lenticellate; mesocarp fleshy; endocarp very hard, fibrous with a 
rough exterior, densely tomentose within, with 2 lateral plates which break away 
on germination. Germination phanerocotylar. Cotyledons fleshy, pale green; 
cataphylls absent; first 2 eophylls opposite, the others alternate or opposite. 

Distr . In tropical Africa 10 species, one native to Central America and one widespread species in Malesia, 
NE. Australia and W. Pacific. 

1. Maranlhes corymbosa Blume, Bijdr. (1825) 89; Craib, F1. Siam. Enum. 1 (1931) 563. - Grymania 

KosTERM. Candollea 20 (1965) 107; Prance & salicifolia Presl, Epim. Bot. (1849) 193; Walp. 

Whitm. Tree Fl. Malaya 2 (1973) 330, excl. syn. Ann. 3 (1853) 854. - Parinarium griffithianum 

Couepia panamensis. - Exitelia corymbosa (Blvme) Be.nth. in Hook., Niger Fl. (1849) 334; Fl. Austr. 2 

Blume, Fl. Java 1, Praef. (1828) vii. - Maranlhes (1864) 426; Walp. Ann. 2 (1851/52) 463; Blume, 

multiflora Korth. Verh. Nat. Ges. Ned. Overz. Mus. Bot. Lugd.-Bat. 2 (1852) 98; Melang. Bot. 2 

Bezitt., Bot. (1839/42) 259; Ned. Kruidk. Arch. 3 (1855) 10; Miq. Fl. Ind. Bat. 1. 1 (1855) 356; ibid. 

(1855) 281; Teijsm. & Bins. Cat. Hort. Bog. (1866) (1858) 1084; Hook. /. Fl. Brit. India 2 (1878) 310; 

253. - Exitelia multiflora (Korth.) Walp. Rep. 5 .Miers, J. Linn. Soc. Bot. 17 (1879) 336; Vidal, 

(1845/46) 115; Miers, J. Linn. Soc. Bot. 17(1879) Sinopsis Atlas (1883) 25; Maingay. Kew Bull. (1890) 

336, sub Exileles. - Parinarium griffithianum 122; King, J. As. Soc. Beng. 66 (1897) 283; Bailey, 

Benth. in Hook.. Niger Fl. (1849)334; Fl. Austr. 2 Queensl. Fl. 2(1900) 524; K. & V. Bijdr. 5 (1900) 334; 

(1864) 426; Walp. Ann. 2 (1851/52) 463; Blume, K.Sch. & Laut. Fl. Deut. Schutzgeb. Siidsee (1901) 

Mus. Bot. Lugd.-Bat. 2 (1852) 98; Melang. Bot. 2 341; Perk. Fragm. Fl. Philip. (1904) 118; Brandis, 

(1855) 10; Mio. Fl. Ind. Bat. I, 1 (1855) 356; ibid. Indian Trees (1906) 278; Foxw. Philip. J. Sc. 2 

(1858) 1084; Hocjk. /. Fl. Brit. India 2 (1878) 310; (1907) Bot. 386; Backer, Schoolfi. Java (191 1) 446; 

Miers, J. Linn. Soc. Bot. 17 (1879) 336; Vidal, Ridley, Fl. Mai. Pen. 1 (1922)670; Disp. (1930)400; 

Sinopsis Atlas (1883) 25; Maingay, Kew Bull. (1890) Craib, Fl. Siam. Enum. 1 (1931) 563. - Parinarium 

122; King. J. As. Soc. Beng. 66(1897) 283; Bailey, maranlhes BiVMt., Mus. Bot. Lugd.-Bat. 2(1852)99; 

Queensl. FI.2(I900)524;K.&V. Bijdr. 5(1900)334; Melang. Bot. 2 (1855) 10. - Parinarium corym- 

K.Sc-H. & Laut. Fl. Deut. Schutzgeb. Sudsec (1901) hosum (Blume) Miq. Fl. Ind. Bat. 1, 1 (1855) 356; 

341; Pf.rk. Fragm. Fl. Philip. (1904) 118; Brandis. ibid. (1858) 1084; Ann. Mus. Bot. Lugd.-Bat. 3 

Indian Trees (1906) 278; Foxw. Philip. J. Sc. 2 (1867) 237; Walp. Ann. 4 (1857)645; Vidal, Cat. PI. 

(1907) Boi, 386; BArKFR. SchoolH. Java (191 1) 446; Silv. Cull. Manila (1880) 29; Merr. Philip. J. 

RirjLF.Y,Fl. Mai, Pen. I (1922)670; Disp. (1930)400; Sc. 10 (1915) Bot. 309; Spec. Blanc. (1918) 162; 


Flora Malesiana 

[ser. I, vol. 10^ 




Fig. 22. Distribution of Maranthes corymbosa Blume. 

Enum. Born. (1921) 290; Enum. Philip. Fl. PI. 2 
(1923) 235; Craib, Fl. Siam. Enum. 1 (1931) 563; 
BuRK. Diet. (1935) 1695; Corner, Wayside Trees 
(1940) 527; Backer & Bakh./. Fl. Java 1 (1964) 522. 
- Parinarium mulliflorum (Korth.) Miq. Fl. Ind. 
Bat. 1, 1 (1855) 356; ibid. (1858) 1084; Suppl. 
Sumatra (1860) 115; ibid. (1861) 307; C.Muell. in 
Walp., Ann. 4 (1857) 646. - Parinarium salicifolium 
(Presl) Miq. Fl. Ind. Bat. 1 , 1 (1855) 357; C.Muell. 
in Walp., Ann. 4 (1857) 646. - Maranthes speciosa 
KoRTH. ex Miq. Fl. Ind. Bat. 1, 1 (1855) 357. - 
Chrysobalanus ciliatus Korth. ex Miq. I.e. 357. - 
Petrocarya griffithiana (Benth.) Miers, J. Linn. 
Soc. Bot. 17 (1879) 336. - Parinarium racemosum 
ViDAL, Cat. PI. Len. Silv. Cult. Manila (1880) 29. - 
Ferolia griffithiana (Benth.) O. Ktze, Rev. Gen. PI. 
1 (1891)216. - Ferolia corymbosa (BtvuE) O. Ktze, 
I.e. 216. - Ferolia salicifolia (Presl) O. Ktze, I.e. 
216. - Parinarium nitidum auct. non Benth.: 
KooRD. Meded. Lands Planten Tuin Btzg 19 (1898) 
448. - Polyalthia pulchrinervia Boerl. Cat. PI. 
Hort. Bog. (1899) 20; Icon. Bog. 1 (1899) 106. - 
Parinarium palauense Kanehira, Bot. Mag. Tokyo 
45 (1931) 282;~ Fl. Micrones. (1933) 129; J. Dept. 
Agr. Kyushu Imp. Univ. Fukuoka 4 (1934) 325. - 
Fig. 21. 

Small to large tree up to 40 m, sometimes flower- 
ing when only a few metres high, trunk not but- 

tressed or slightly enlarged at base. Stipules intrapet- 
iolar, lanceolate, acute, 5-10 mm long, sparsely 
pilose on exterior, glabrous within, early deciduous. 
Leaves coriaceous, usually oblong-lanceolate to 
oblong-elliptic, 6.5-14 by 2.5-8 cm, acuminate at 
apex, the acumen 8-20(-30) mm long, cuneate at 
base, glabrous when mature but often sparsely cadu- 
cous arachnoid-lanate when young, usually with 2 
conspicuous prominent glands at junction of petiole 
and decurrent lower surface; primary veins 7-10 
pairs, arcuate, prominulous on both surfaces; midrib 
plane above, prominulous beneath; petioles 4-9 mm 
long, glabrous when mature, flattened above. In- 
florescences of flattened many-flowered corymbose 
panicles, rachis and branches sparsely pilose, 
glabrescent. Bracts and bracteoles ovate to lanceo- 
late, sparsely pubescent, caducous. Receptacle tur- 
binate, tapering into pedicels 2-4 mm long, grey 
tomentose to glabrous on exterior, glabrous within, 
calyx lobes fleshy, ovate to elliptic, obtuse, 2.5-4 
mm long, unequal. Petals white tinged pink, gla- 
brous, 3-6 mm long, caducous. Stamens 25-35 in- 
serted in several rows on one side of throat, with 
tooth-like staminodes opposite. Ovary bilocular, 
densely lanate and villous. Style glabrous except at 
base; stigma truncate. Fruit ellipsoid, 3-4 mm long, 
1 .5-2 cm broad, tapered towards base; epicarp thin, 
glabrous on exterior when mature, sometimes lanate 


Chrysobalanaceae (Prance) 


when young; endocarp hard, 5 mm thick, rough on 
exterior; densely lanate within; bilocular usually with 
seed in one locule only. Cotyledons plane-convex. 

Disir. S. Thailand extending east to Solomon 
and Caroline Islands and Australia (Queensland, 
Northern Territory); in Malesia: Malay Peninsula, 
Sumatra, Java, Borneo, Lesser Sunda Islands, 
Sulawesi, Philippines, Moluccas, New Guinea, New 
Britain and Admiralty Islands. Fig. 22. 

Ecol. Common in coastal areas on rocky and 
sandy hills and extremely inland up to 600 m altitude. 
Also in gallery forest and in Australia on sand dunes 
behind mangrove swamp. In Kalimantan the fruit is 
eaten by many bird species, including hornbills and 
fruit pigeons, which probably disperse the seed. The 
seed is also scatter-hoarded by the squirrel Sun- 
dasciurus hippurus. African species of Maranthes 
are bat-pollinated. 

Uses. Wood used for house-building and for 
posts. Fruit edible. 

Vern. Thailand: chi-kai-pen, chi-ot-pen, Korat; 
Malay Peninsula: chana, lejin, merbatu, m. layang, 
mujagon, sau hulan, sunko rimau; Sumatra: damor 
lilis, kajiebatu, kaju baiu, Banka, kalek kureseng, k. 
parada; Java: gesing, kituwat, solo, sulo, triwulan, 
Huloh, Jav., taritik, l. monjet, Sund.; Borneo: bang- 

kawang, bonsissian, Malay; bansisian, Sabah, 
Tengara; nyalin laat, Sarawak; buenza, kajebabu, 
kajoe kambang, kambang, potang, Kalimantan; 
Sulawesi: kolaka; Tidore: latan, Aru Is.; Philip- 
pines: almag, delebaybai, kaphangan, kolaka, kola- 
sa, kulingan, malapiga, malapuyan, sampinit, tak- 
dangan. Tag., aningat, binggas, caratacat, 
kagemkena, karatakat. Ilk., arangan, Tagb., daka- 
yau. Pang., bakoyan, tapas, P. Bis., bongog, dau, 
mata-mata, sarangun, S.-L.Bis., dumaga, Kuy., ka- 
gangan, kalakangon, ogat. Bag., kamuli tingan, 
Pamp., lank angan, Lan., langog, Buk., lumaluas, 
sigaadan, Mag., maluktik, Sul., salipungan, salutui, 
Neg., bareraga, baril, Bik., C.Bis., laiusin, Bik., 
S.C.Bis., liusin, Sbl., Tag., Bik., sabongkaag. Ilk., 
Ting., tadiang manok. Ting., Tag.; New Guinea: 
badigal, Wagu, djuramun, Kemtuk, jambuan, 
Kaigorin, kaupen, Jal, kawol, kowot, Muyu, kwanu, 
Maprik, lakan, luikoko. Bush Mekeo, marigag, 
Sinai, mehlue, Bembi, morolee, mun, Dagu, naas, 
ningua, njali, Nemo, njiwa, niwa, Sidei, paguh, 
Timbunke, phu, Wasuk, watu, Karopa; Solomon 
Is.: asikisiki, giza, mon warlu, morigag, now-wa-ru, 
santalan; Bougainville: mon-warku, Kugumaru, 
marigai, Siwai, Bouin; Palau Is.: apgau. 


Prance [Tree FI. Malaya 2 (1973) 327, unpublished], Brittonia 31 (1979) 91; 
Prance & White, Phil. Trans. Roy. Soc. Lond. 320 (1988) 149, f. 40, 41. - 
Parinari auct. non Aubl.: quoad P. heteropetala Scortech. ex King et P. 
myriandra Merr., tan turn. - Acioa auct. non Aubl.: Kosterm. Reinwardtia 
7 (1965) 9. - Fig. 23. 

Large trees, ultimate shoots not divaricate. Stipules to 7 mm long, foliaceous, 
persistent, lanceolate to ovate. Leaves glabrous on both surfaces with minute 
papillae on veins giving a beaded appearance. Petioles eglandular. Inflorescence 
an unbranched or little-branched terminal or axillary raceme with shortly 
stalked congested cymules proximally and singly inserted flowers distally. 
Bracts and bracteoles small, suborbicular, persistent, eglandular, not enclosing 
groups of flower buds. Flowers hermaphrodite, strongly zygomorphic. Recep- 
tacle broadly obconic-campanulate, shorter than calyx lobes, asymmetric, 
hollow, hairy on both surfaces, but throat not blocked by retrorse hairs; calyx 
lobes 5, markedly unequal, suborbicular to lingulate, strongly imbricate. Petals 
5, unequal in size and shape, the 2 posterior larger than the others, markedly 
ungulate and enclosing stamens in bud. Stamens 8-30, inserted unilaterally on 
margin of disk; filaments united for half to three quarters of length into a strap; 
sta.ninodcs 5 8, inserted opposite stamens. Ovary inserted laterally at mouth 
of receptacle; unilocular with 2 ovules. Fruit large, hard; epicarp glabrous, 


Flora Malesiana 

[ser. I, vol. 10^ 

f D K.TincL 

Fig 23. Kostermanthus heteropetalus (Scortech. ex King) Prance. A. Habit; ^•^"°^^^^^f^'°"' f„f ^ 
petals; £. stamen; F. fruit; C. ovary section (>»-£, G Ogata KEP 105153, FMeijer SAN 34279). 


Chrysobalanaceae (Prance) 


crustaceous-verrucose; endocarp hard, thick, glabrous within, breaking ir- 
regularly on germination. Cotyledons slightly ruminate. 

Distr. Malesia: Malay Peninsula, Sumatra, Borneo, Sulawesi, Philippines (Mindanao); 2 species. 


1. Leaves coriaceous; petioles 6-12 mm long; calyx tube 2-3 mm long, broadly campanulate 

1. K. heteropetalus 
1 . Leaves chartaceous; petioles 2-3 mm long; calyx tube 5 mm long, slender 2. K. malayanus 

1. Kostermantbus heteropetalus (Scortech. ex 
King) Prance, Brittonia 31 (1979) 91; Prance & 
White, Phil. Trans. Roy. Soc. Lond. 320 (1988) 152 
- Parinarium heteropetalum Scortech. ex King, J. 
As. Soc. Beng. 66 (1897) 283; Ridley, F1. Mai. Pen. 
1 (1922) 670; Nayaranaswami, J. As. Soc. Beng. 
n.s. 27 (1931) 368. - Parinarium kunstleri King, J. 
As. Soc. Beng. 66 (1897) 282; Ridley, F1. Mai. Pen. 
1 (1922) 670. - Parinarium myriandrum Merr. 
Univ. Cal. Publ. Bot. 15 (1929) 93. - Acioa hetero- 
peiala (Scortech. ex King) Koster.m. Reinwardtia 7 
(1%5) 11. - Fig. 23. 

Tree to 35 m tall, older trees buttressed to 1 m up 
trunk; young branches glabrous, lenticellate. Stip- 
ules 6-7 mm long, partly intrapetiolar, carinate, 
ovate, foliaceous, acute to acuminate, persistent to 
subpersistent. Leaves coriaceous, usually elliptic- 
subovate to rarely lanceolate, 5-20 by 2.5-6 cm, 
bluntly acuminate at apex, cuneate to rounded at 
base, glabrous on both surfaces, minutely papillose 
on venation of both surfaces giving a bead-like ap- 
pearance; midrib prominulous above, prominent 
beneath; primary veins 6-10 pairs, arcuate, slender, 
prominent beneath; petioles 6-12 mm long, some- 
times lightly alate from decurrent leaf margins, 
slightly flattened above, eglandular. Inflorescences 
little-branched, to 10 cm long, the rachis and branch- 
es lightly tomentellous; bracts and bracteoles ovate, 
acute, to 3 mm long, caducous. Receptacle broadly 
campanulate, 2-3 mm long, tomentose on both sur- 
faces; calyx lobes fleshy, unequal, acute, to 7 mm 
long, pilose on both surfaces, reflexed in open flow- 
ers. Petals white tinged pink, fleshy, ellliptic, con- 
cave, largest up to 15 mm long, tomentellous on ex- 
terior, enveloping staminal ligule, the others much 
smaller to 6 mm long. Stamens 25-30 united into a 
unilateral ligule for 2/3 length, to 12 mm long, 
glabrous; anthers pubescent. Ovary densely pilose. 
.S7>'/^ densely appressed pilose, stigma truncate. Fruit 
ovoid, unilocular 4 by 3 cm; epicarp glabrous, crus- 
taceous; endocarp hard, thick. Cotyledons slightly 
ruminate, 1.5 by 3 cm. 

Distr. Malesia: Malay Peninsula, Sumatra, 
Borneo, Sulawesi, Philippines (Mindanao). Fig. 

Ecol . From sea level up to 500 m altitude. 

Fig. 24. Distribution of Kostermanthus heterope- 
talus (Scortech. ex King) Prance (dots) and K. 
malayanus (Kosterm.) Prance (star). 

2. Kostermanthus malayanus (Kosterm.) Prance, 
Brittonia 31 (1979) 94; Prance & White, Phil. 
Trans. Roy. Soc. Lond. 320(1988) 152. - Acioa ma- 
layana Kosterm. Reinwardtia 7 (1965) 13. 

Small tree to 10 m; young branches glabrous, len- 
ticellate. Stipules lanceolate, acute, glabrous, sub- 
persistent, c. 5 mm long. Leaves chartaceous, ellip- 
tic, 14-20 by 6.5-8.5 cm, acuminate at apex, the 
acumen 4-10 mm long, cuneate at base, glabrous on 
both surfaces; midrib slightly prominulous to plane 
above, prominent beneath, with a pair of round 
glands at base; primary veins 10-13 pairs, pro- 
minulous above, prominent beneath; petioles 2-3 
mm long, glabrous, slightly alate with decurrent leaf 
margins. Inflorescence of subterminal racemes or lit- 
tle branched, the rachis brown pilose pubescent; 
bracts and bracteoles ovate, acute, to 3 mm long, 
caducous. /?ecfp/flc/e slender cylindrical, 5 mm long, 
sessile tomentose on exterior, densely tomentose 
within; calyx lobes ovate, acute, 4-5 mm long, 
densely tomentose on exterior, glabrous within ex- 
cept at apex. Petals spalhulate, 6 mm long, clawed. 
Stamens S- 10, united into a unilateral ligule for half 
of length. Ovary densely pilose. Style pilose for most 
of length. Fruit unknown. 

Distr. Malesia: Malay Peninsula (Penang). 
Known only from the type collection. 


Flora Malesiana [ser. I, vol. 10^ 

Insufficiently known 

Acioa percohacea Kosterm. Reinwardtia 7 (1965) 14. j j- . •• u ^ «• 

This species was described from a single sterile collection from the Malay Penmsula, and distmgu.shed from 

Kostermanthus heteropetalus Prance by its pubescent branches and caducous pubescent leaf undersurfaces. 

It is impossible to evaluate until further material is collected, but almost certamly belongs withm K. 


SABIACEAE (C.F. van Beusekom & Th.P.M. van de Water, Leiden)* 

Trees, scandent shrubs or woody climbers. Leaves alternate or spirally ar- 
ranged, penninerved, simple or imparipinnate, the leaflets in the latter case op- 
posite on often somewhat swollen nodes of the rachis; exstipulate. Flowers 
small, bisexual, rarely polygamo-dioecious, in terminal or axillary racemose 
panicles, or cymose: paniculately arranged cymes, or these reduced to solitary 
axillary flowers. Sepals (3-)5, imbricate, free or ± connate at the base, equal 
or unequal. Petals (4-)5, mostly opposite the sepals (rarely alternate: 
Ophiocaryon spp. , South America). Stamens (including staminodes) 5, opposite 
the petals, all polliniferous (Sabia) or only 2 inner ones opposite the reduced 
petals polliniferous and the other 3 staminodial. Disk small, annular, surround- 
ing the base of the ovary. Ovary of 2(-3) carpels united to form a compound 
superior ovary, carpels very rarely free in the apical part, in that case tapering 
to 3 short styles with a capitate stigma; otherwise normally a short, cylindric or 
conical style; cells 2(-3), each with 1 or 2 pendulous or horizontal, axile hemi- 
tropous, unitegmic, crassinucellar ovules. Fruit either 1 -celled or 2-coccous, 
drupaceous or dry, indehiscent; endocarp often wrinkled. Endosperm scanty or 
wanting. Embryo with a curved radicle and 2 folded or coiled cotyledons. 

Distribution. Three genera: Sabia Indo-Malesian, from the S. Deccan and Kashmir to S. 
Japan, throughout Malesia as far as the Solomons; Meliosma with a similar range but also occur- 
ring in tropical America; Ophiocaryon in the Neotropics. The family is absent in Australia and 

Fossils of both Malesian genera are found onwards of the Oligocene and Eocene in Asia and 
Europe. See under the genera. 

Ecology. Tropical forests, mostly below 2000 m altitude. 

Taxonomy & Delimitation. There is no concensus of opinion on the affinity, hence the 
systematic position of Sabiaceae. Some even doubt whether Sabia and Meliosma are correctly 
placed in one family. 

After the description of Sabia by Colebrooke (1818), Blume (1851) accommodated it in a new 
monogeneric family, Sabiaceae, suggesting its affinity with Menispermaceae. Shortly afterwards 
MiERS (see Lindley, 1853), while working on Menispermaceae, placed Sabia between that family 
and Lardizabalaceae . Hooker/. & Thomson (1855) considered the genus intermediate between 
Menispermaceae and Schisandraceae. 

The scandent habit and the resemblance of the drupelets of Sabia with those of Menispermaceae 
undoubtedly were a major argument for supposed affinity. 

Subsequently Bentham & Hooker (1862) extended the then monogeneric family Sabiaceae to 
include Meliosmaceae Endl., adding the genera Meliosma Blume and Ophiocaryon Schomb.; 
both are trees, the first Asian-American, the latter tropical American. They removed the family 
in its new concept from the Menispermaceous affinity and accommodated Sabiaceae near Sapin- 
daceae and Anacardiaceae. This position has been stable for a century and was adhered to by 
many leading botanists: Warburg (1895), von Wettstein (1911), Hutchinson (1926, 1973), 
Melchior (1964). Takhtajan (1969), Dahloren (1975, 1983), and Thorne (1976, 1983). Some 
of these authors showed some doubt about the position and some made suggestions, e.g. War- 
burg {I.e. 370), who believed one could possibly derive the flower of Meliosma from the Meni- 

( I ) Accommodated from t he monographs of both authors in Biumea volumes 1 9 and 26, and provided with 
an introduction. 



Flora Malesiana [ser. I, vol. 10^ 

spermaceous scheme and mentioned that Radlkofer was not in favour of an affinity with Sapin- 
daceae or Anacardiaceae. 

In recent years there is a tendency to return to Blume's opinion towards affinity with Menisper- 
maceae. Pollen morphology (Erdtman, 1952) and embryology (Mauritzon, 1936) have been in- 
terpreted in favour of a relationship with Menispermaceae. Airy Shaw (1973) remarked that the 
opposition of calyx, corolla and stamens is a most unusual feature, but can probably be derived 
from the Menispermaceous type of fiower. In his recent classification Cronquist (1981) tentative- 
ly placed Sabiaceae near Menispermaceae in the Ranunculales. Also Forman, in his treatment of 
the Menispermaceae (Fl. Males. I, 10^ 1986, 157-253), shares this opinion. 

Another matter is whether Sabia and Meliosma/Ophiocaryon should be accommodated in one 
family; hitherto they are represented by two tribes in Sabiaceae (Warburg, 1890), differing in 
habit (climbers versus trees), the leaves, and in the androecium. Moreover, Cronquist (1981) 
mentioned in his discussion that, according to Wolfe, the leaf venation of Sabia is highly com- 
patible with a position near Menispermaceae, but that of Meliosma more similar with some 
members of the Rosidae. There may be more arguments to accommodate Meliosma in a separate 
family Meliosmaceae Endl., apart from Sabiaceae sensu stricto. This opinion was held by Airy 
Shaw (1973). 

References: Airy Shaw in Willis, Diet. ed. 8 (1973) 1017; Bentham & Hooker, Genera Plan- 
tarum 1 (1862)413; Blume, Mus. Bot. Lugd. -Bat. 1 (1851) 369; Cronquist, An integrated system 
of classification of flowering plants (1981) 140; Dahlgren, Bot. Notis. 128 (1975) 126; Nordic 
J. Bot. 3 (1983) 144; Erdtman, Pollen morphology and plant taxonomy (1952) 380; Hooker/. 
& Thomson, Flora Indica 1 (1855) 208; Hutchinson, Families of flowering plants 1 (1926) 254; 
ed. 3 (1973) 449; Lindley, Vegetable kingdom ed. 3 (1853) 467; Mauritzon, Acta Hort. Goth. 
1 1 (1936) 18; Melchior, Engler's Syllabus 2 (1964) 285; Takhtajan, Flowering plants: origin and 
dispersal (1969) 226; Thorne, Evol. Biol. 9 (1976) 61 ; Nordic J. Bot. 3 (1983) 106; Ware, in E. & 
P., Nat. Pfl. Fam. 3, 5 (1895) 367; Wettstein, Handb. Syst. Bot. ed. 2 (1911) 633. 

Vegetative Anatomy. - Leaf anatomy. Hairs unicellular in Saft/o; uniseriate nonglandular 
and capitate glandular in Meliosma. Stomata confined to the lower leaf surface, anomocytic or 
paracytic. Mesophyll dorsiventral, with arm palisade cells in Meliosma. Veins embedded in 
mesophyll and sheathed by sclerenchyma. Petiole in distal end with a closed vascular cylinder. 
Crystalliferous cells containing clusters common near the veins. 

Young stems. Cork superficial. Cortex with stone cells in some species oi Meliosma. Pericyclic 
sclerenchyma forming a composite, closed ring in Sabia, and composed of isolated fibre groups 
in Meliosma. Phloem with broad lignified rays in Sabia, and with non-lignified, dilatating (tri- 
angular) rays in Meliosma. Vessels with mixed simple and scalariform perforations in first formed 
xylem. Cluster crystals common in cortex, phloem, and pith. Secretory cells with unidentified con- 
tents noted in parenchyma of several Meliosma species. 

Wood anatomy. Vessels exclusively solitary in Sabia, solitary and in radial multiples or small 
clusters in Meliosma; vessel perforations typically simple in Sabia; mixed simple and scalariform 
or exclusively scalariform to reticulate in Meliosma. Intervessel pits alternate. Vessel— ray and 
vessel-parenchyma pits simple, and often large. Fibres, usually thin-walled, with minutely 
bordered to simple pits, and mainly confined to the radial walls in Meliosma (libriform fibres); 
with distinctly bordered pits common in both the radial and tangential walls in Sabia; occasionally 
septate. Parenchyma scanty paratracheal to vasicentric with occasional lateral extensions in 
Meliosma, very sparse to almost absent in Sabia, usually in 8-celled strands. Rays sometimes of 
two different sizes, the broad ones 4-8(- 15) cells wide in Meliosma, up to 20 cells wide in Sabia, 
usually over 2 mm high, heterogeneous (Kribs type II), often with sheath cells. 

Taxonomic note based on vegetative anatomy. The above description is mainly based on early 
studies of a very limited number of species, so that the information is far too limited to serve in 
the discussion of infrageneric classification and delimitation. The two genera are anatomically 
quite distinct in their leaf and wood anatomy. Partly this is related to general anatomical dif- 

1989] Sabiaceae (van Beusekom & van de Water) 681 

ferences between climbers (Sabia) and erect shrubs or trees (Meliosma). Thus, the anatomical 
evidence can be interpreted both in favour of the separation of Meliosma and Sabia into two 
families, or alternatively to retain their tribal position in the same family. Anatomically Sabia is 
quite distinct from the Menispermaceae to which it has been compared (see above, under tax- 
onomy); affinity oi Meliosma and Sabia with families of the Sapindales, especially A nacardiaceae 
seem to find more support in vegetative anatomy. 

References: Carlqvist, Aliso 11 (1985) 139-157; Desch, Manualof Malayan Timbers 2 (1954) 
522-523; Metcalfe & Chalk, Anatomy of the Dicotyledons 1 (1950) 448-452; Moll & 
Janssonius, Mikrographie des Holzes 2 (1922) 424-437; Solereder, Systematische Anatomic 
der Dicotyledonen (1899) 276-278; & Erganzungsband (1908) 108-109. - P. Baas. 

Palynology. Pollen grains in Sabiaceae are prolate spheroidal to prolate. Size ranges from 
20 to 33 |im. The apertural system is always tricolporate. Ectoapertures are long colpi, endoaper- 
tures are lalongate pori or short colpi. The shape of the endoapertures is oblong to elliptic, 
sometimes appro.ximately rectangular or meridionally constricted. Exine stratification is easily to 
observe in the light microscope. Each layer is about uniformly thick throughout. The tectum is 
equally thick or up to twice as thick as the nexine. It is mostly more than twice as thick as the 
columellate layer. Total exine thickness is 1-2.5 |im. The ornamentation is usually finely to 
coarsely reticulate; sometimes it is finely or indistinctly perforate. 

Meliosma and Sabia show only little infrageneric variation. Moreover, the ranges in both 
genera are rather similar. Only minor differences exist: Sabia mostly has a thinner exine with a 
finer reticulate ornamentation than Meliosma. Pollen morphology does not support accom- 
modating the genera in separate families (Mondal & Mitra, 1982). 

As taxonomists, pollen morphologists are ambiguous with respect to the position of the 
Sabiaceae. Erdtman (1952) reported pollen similar to that of Sabiaceae to occur in several other 
families. However, heactually mentioned only the A/e/?/5/?ermcrceae. PoUen of A nacardiaceae and 
Sapindaceae was considered less similar or different. According to Mondal & Mitra (I.e.) 
Sabiaceae pollen differs from that of Aceraceae, Hippocastanaceae, Lardizabalaceae, Melian- 
thaceae, Menispermaceae, Sapindaceae, and Schizandraceae. On the basis of grain shape and 
size, P/E ratio, exine structure and aperture characters they suggested to classify the Sabiaceae 
nearest to the Anacardiaceae. It must be stressed, however, that it is extremely difficult to infer 
relationships from resemblances between rather simple pollen types. Obviously unrelated taxa 
may show very similar pollen, whereas closely related taxa sometimes have completely different 

References: Erdtman, Pollen morphology and plant taxonomy, Angiosperms (1952) 390; 
Mondal & Mitra, Geophytology 12 (1982) 166-180. - R.W.J.M. van der Ham. 

Phytochemistry. The only observations worth to be reported here are the presence of pen- 
tacyclic triterpenoids of the oleanene series and the absence of starch in seeds. The 3-acetates of 
oleanolic acid and oleanolic aldehyde were isolated from bark of Meliosma simplicifolia. Seeds 
of Meliosma myriantha Sieb. & Zvcc. (continental SE. Asia) were reported to give positive reac- 
tions for alkaloids and to contain 8% of protein and 10<^o of fatty oil but no starch. 

References: Desai c.s., Indian J. Chem. 15B (1977) 291; Hegnauer, Chemotaxonomie der 
Pnanzen 6 (1973) 240. - R. Hegnauer. 

Note. Though the genera are extremely clearly defined, specific delimitation has in both 
genera been difficult, as it seems that racial segregation is common in both. Van de Water has 
in Sabia employed a finer specific distinction than van Beusekom did in Meliosma. 

KEY TO the genera 

I . Climbers or scandent shrubs. Flowers with 5 equal, fertile stamens, in usually rather few-flowered thyrses 
or cymes, sometimes reduced to a single axillary flower. Leaves simple, entire or subcntirc, alternate 

I. Sabia 

682 Flora Malesiana [ser. I, vol. 10"^ 

1. Trees. Flowers in usually large, racemose, terminal or axillary panicles. Fertile stamens only 2, the other 
3 abortive and reduced to scales or nectary-like bodies. Leaves uneven pinnate, leaflets opposite on ± nod- 
ed rachis, rarely simple, entire or toothed, spirally arranged 2. Meliosma 


CoLEBROOKE, Trans. Linn. Soc. Lond. 12 (1818) 355, t. 14; Wall, in Roxb., Fl. 
Ind. 2 (1824) 308; Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 368; Ware, in E. & 
P., Nat. Pn. Fam. 3, 5 (1895) 367, f. 183A, 184A-H; Chen, Sargentia 3 (1943) 
1; VAN DE Water, Blumea 26 (1980) 1. - Meniscosta Blume, Bijdr. (1825) 28; 
DiETR. Syn. PI. 2 (1840) 923 ('Menicosta'). - Fig. 2-4. 

Evergreen or deciduous, woody climbers or more or less scandent shrubs 
(rarely recorded as small trees). Twigs terete, striate (see note), with ± promi- 
nent leaf cushions, unarmed, mainly in deciduous species with some cataphylls 
at their base, spirally arranged. Buds either ± globular and obtuse to rounded, 
or ovoid and acute; scales glabrous to pubescent, ciliolate or not, persistent at 
the base of the twigs. Leaves simple, ovate or elliptic to lanceolate, 2-25 by 
1-10 cm, herbaceous to coriaceous, petioled, entire or very rarely subentire; 
nerves 3-12 pairs, ascending to patent, curved to straight. Flowers bisexual, 5- 
merous, actinomorphic, up to c. 15 mm diam., green to white, yellow, or pur- 
ple, axillary, either solitary, or arranged in a few- to many-flowered cyme, ap- 
pearing before or with the new leaves. Cymes axillary, either solitary, or, when 
the subtending leaves are shed or are bract-like, arranged in racemose to thyr- 
soid or sometimes corymbose inflorescence, pedicel ± thickened upwards in 
fruit; bracts ovate to lanceolate, up to 6 mm, bracteoles as bracts but usually 
smaller, or sepal-like, or minute and then often situated near calyx. Sepals 5(-7, 
see bracteoles), equal to very unequal mutually, mostly ± confluent at the base, 
variable in size and shape but often suborbicular or broad-ovate to ovate, persis- 
tent. Petals 5, rarely 6 or 7, episepalous, imbricate, suborbicular to lanceolate, 
glabrous, sometimes (sub)ciliolate, persistent or not; nerves ± parallel, branch- 
ing or not, sometimes conspicuous when dark-coloured. Stamens 5, epipeta- 
lous, ± equal, persistent or not; filaments more or less flattened, adherent to 
the base of the subtending petals; anthers globular to ellipsoid, introrse, upright 
or inflexed. Disk in most species ± crown-shaped, sometimes short-cylindrical 
{S. sumatrana), truncated conical, or ± cushion-shaped; lobes and ribs, if pres- 
ent, alternating with the stamens. Pistil: style conical to cylindrical, rarely ab- 
sent, persistent. Ovary superior, 2-celled, (sub)globose to subreniform, usually 
laterally somewhat compressed, very rarely subapocarpous. Ovules 2 per cell, 
more or less superimposed, attached to the septum, hemi-anatropous. 'Drupe- 
lets' 1-seeded or very rarely with 2 seeds, (sub)globose, obovoid, oblong- 
obovoid (or pyriform), or subreniform, laterally ± compressed, green or white 
to red or deep blue when fresh; mesocarp rather thin, pulpy, sometimes with 
many dark 'granules', endocarp crustaceous, very often with ± prominent ribs 


Sabiaceae (van Beusekom & van de Water) 


Fig. 1 . The Southeast Asian and Malesian distribution area of Sabia Colebrooke. The numbers refer to the 

number of species in that area. 

forming a fine to coarse reticulate pattern, margin sometimes distinctly keeled. 
Seed conform to the drupelet; testa usually conspicuously dark-dotted, inside 
often lined with a very thin layer of endosperm. Embryo with two flat, smooth, 
somewhat undulated, or sometimes strongly folded cotyledons and a cylindrical 
rootlet curving to the hilum. 

Distr. Indo-Malesia, along the Himalayas (1 species disjunct, also in the S. Deccan) through Burma and 
China to S. Japan; throughout Malesia (not yet known from the Lesser Sunda Islands), as far as New Guinea, 
the Louisiades and Solomon Islands. In all 19 species, of which 7 in Malesia. Fig. 1. 

Ecol. Inconspicuous climbers (rarely reported as small trees), except two continental Asian species all 
evergreen, found in forests and thickets, from the lowland up to c. 1000-1200 m altitude, 5. javanica up to 
1500 m and 5. pauciflora to 2000 m; S. racemosa ssp. kinabaluensis is mainly montane, at 800-1500 m. 
Flowering occurs mostly throughout the year. 


I . Flowers solitary, sometimes 2 or 3 together, or arranged in a thyrsus; ovary glabrous; style in flower 3-6 
mm long, conspicuous in fruit and about half as long as the adjacent side(s) of the drupelet(s) 

7. S. sumatrana 
1 . Flowers in cymes, these solitary, axillary, up to 30(-40)-nowered; ovary densely pubescent; style in flower 

2.25-2.5 mm long; drupelets not known 1. S. erratica 

I Flowers in few- to many-flowered cymes; cymes either solitary, axillary, or arranged in an up to 15 cm long 

racemose to thyrsoid inflorescence, (I - )2-25-nowercd; ovary glabrous; style in flower up to 1.5(-1.75) 

mm long, inconspicuous in fruit and much shorter than the adjacent side(s) of the drupelct(s). 

2. Leaves oblong to lanceolate, 3- I2(- 15) by 1-5 cm, beneath usually distinctly paler than above; nerves 

(5-)6-9(-IO) pairs, patent, straight; cymes solitary, axillary, (4-)7-25-nowered; style either absent or 

obscurely or normally developed, (0.75-) I - 1 .5(- 1 .75) mm long 4. S. parviflora 

2. Leaves elliptic-oblong to sublanccolatc, 5-25 by 2- 10 cm, beneath usually somewhat paler than above 
but not conspicuously so; nerves 4 8(-9) pairs, ± patent, straight to curved; cymes often arranged in 
an up to 15 cm long racemose to thyrsoid inflorescence, (l-)2- I0( l2)-nowercd, sometimes solitary, ax- 
illary, up to 4( 6)-nowcred; style normal-developed, 0.2- I mm long. 
3. Cymes up to 2 cm, 1 4( 6)-nowcrcd; petals suborbicular to elliptic, 1.75 2.5 by 1.25-2 mm, obtuse 


Flora Malesiana 

[ser. I, vol. 10^ 

to rounded; stamens nearly as long as petals; drupelets globular to obovoid, very compressed, 1 1 - 14 by 

10-13 mm; reticulate pattern usually faint or absent 3. S. limoniacea 

3. Cymes up to 1 cm, l-4(-7)-nowered; petals either oblong-ovate to ovate-lanceolate, acute, acuminate 
or not, or elliptic-oblong to oblong, obtuse, 3.5-6.5 by (1 .25-)l .5-2.5 mm; stamens distinctly shorter 
than petals; drupelets obovoid, ± compressed, c. 10-12by(7-)8- 10 mm; reticulate pattern rather faint 

but usually visible, often limited to the margin 6. S. racemosa 

3. Cymes up to 2(-3. 5) cm, (l-)2-10(-12)-flowered; petals oblong, 2.5-4(-4.5) by c. 1-1.5 mm, obtuse; 
stamens distinctly shorter than petals; drupelets obovoid or ± globular, ± compressed, 7.5-11 by 
8-10(-ll) mm; reticulate pattern usually clearly visible, sometimes obscure, limited to the margin or 
4. Leaves oblong to sublanceolate, 5- 14(- 1 8) by 2-6(-8) cm; nerves (5-)6-8(-9) pairs; cymes either ar- 
ranged in a racemose to thyrsoid inflorescence, or solitary, axillary, 1-4-flowered; style 0.6-1 mm; 
drupelets ± globular, sometimes somewhat obovoid, compressed, 7.5-11 by 8-10(-ll) mm 

5. S. pauciflora 
4. Leaves elliptic-oblong to oblong, sometimes sublanceolate, 6-19 by 2-8(-10) cm; nerves 4-7(-8) 
pairs; cymes usually arranged in a thyrsoid inflorescence, sometimes subtended by small leaves, 
3- 10(- 12)- flowered; style 0.2-0.5 mm; drupelets obovoid, sometimes globular, somewhat com- 
pressed, 9-11 by c. 9-10 mm 2. S. javanica 

1. Sabia erratica van de Water, Blumea 26 (1980) 

Evergreen, woody. Twigs glabrous to somewhat 
pubescent; flowering twigs up to 2.5 mm diam., ± 
lax-pubescent. Buds ovoid, acute; scales ± pubes- 
cent, ciliolate. Leaver oblong, 5-8 by 2.5-3 cm, in- 
dex 2-2.7, pergamentaceous, above glabrous or still 
sparsely pubescent especially at the base and on 
midrib, beneath laxly pubescent especially on midrib 
and nerves; base acute, apex acute or short-acumi- 
nate; nerves 6-7 pairs, patent, ± .straight to some- 
what curved; petiole up to 1 .5 cm, glabrous to pubes- 
cent. Cymes solitary, axillary up to 4.5 cm, up to 
40-flowered, ± lax-pubescent; pedicels up to 4 mm; 
bracteoles oblong to oblong-ovate, up to 0.8 mm, 
pubescent, ciliolate. Sepals ovate to somewhat ellip- 
tic, 0.8-1 by 0.5-0.75 mm, obtuse to acute, ± 
pubescent, ciliolate. Petals oblong or oblong-ovate 
to sublanceolate or ovate-lanceolate, 3.75-4 by 
1-1.5 mm, acute to narrow-obtuse, subciliolate, 
nerves up to 6, dark-coloured. Stamens 2.3-3 mm; 
filament flattened, 1 .8-2.6 by 0.25-0.4 mm; anther 
ellipsoid to oblong-ellipsoid, c. 0.4-0.6 mm, 
upright. Disk crown-shaped; lobes very short or ab- 
sent; ribs ± prominent. Pistil 2.75-3 mm; style 
narrowly-conical to cylindrical, 2.25-2.5 mm, with 
some hairs at the base; ovary somewhat globular to 
subreniform, 0.5-0.6 by 0.6-0.8 mm, densely 
pubescent. Drupelets not available. 

Distr . Malesia: Singapore (Bt. Timah Res.), only 
known from the type, collected in 1940. 

Notes. In habit somewhat resembling S. par- 
viflora but readily distinguished by floral characters. 

On the label noted as a 'tree, 100 ft', but this is 
suspected to be a wrong annotation or field observa- 
tion or a wrong label. 

2. Sabia javanica (Blume) Backer ex Chen, Sargen- 
tia 3 (1943) 59; Backer & Bakh./. F1. Java 2 (1965) 
144; VAN DE Water, Blumea 26 (1980) 39. - 
Meniscosta javanica Blume, Bijdr. (1825) 29. - 
Meniscosta scandens Blume ex Spreng. Syst. Veg. 4, 

2 (1827) 114, nom. illeg.; Dietr. Syn. PI. 2 (1840) 
923. - Sabia meniscosta Blume, Mus. Bot. Lugd.- 
Bat. 1 (1851) 369, f. 44, nom. illeg., incl. var. firma 
Blume, var. latifolia Blume et var. glabriuscula 
Blume; Miq. F1. Ind. Bat. 1, 2 (1859) 6 1 8 ( 'me«/co5- 
tay, Fl. Arch. Ind. (1870) 71; //j/c^. (1871) pi. 31, /nc/. 
var. elliptica Miq.; Hook./. Fl. Brit. India 2 (1876) 

3 ('menescortay. Backer, Schoolfl. Java (1911) 273; 
KooRD. Exk. Fl. Java 2 (1912) 544. - Sabia elliptica 
(Miq.) Miq. Sum. (1861) 203, 521. - Sabia javanica 
^lume) Chen var. glabriuscula (Blume) Chen, 
Sargentia 3 (1943)61. 

Evergreen woody climber or scandent shrub, up to 
10 m. Twigs glabrous; flowering twigs up to 5 mm 
diam., glabrous or ± pubescent. Buds ovoid, up to 

2 mm, acute; scales glabrous or with few hairs, ± 
ciliolate. Leaves elliptic-oblong to sublanceolate, 
6-19 by 2-8(-10) cm, index 2-3(-4), pergamen- 
taceous to pergamentaceous-coriaceous, above and 
beneath glabrous or with some hairs on midrib; base 
acute to rounded, apex acute, acuminate; nerves 
4-7(-8) pairs, patent, curved to straight; petiole up 
to 2.5 cm, glabrous to sparsely pubescent, ± (fine-) 
wrinkled. Cymes arranged in an axillary, up to 12 cm 
long, glabrous to pubescent, thryrsoid inflorescence, 
subtended by bracts or sometimes by small leaves 
and then inflorescence up to 17 cm long; cymes up to 

3 cm, forming a lax to dense cluster of 3-10(-12) 
flowers, subglabrous to pubescent. Bracts ovate to 
sublanceolate, up to 5 mm, subglabrous to more or 
less pubescent, ± ciliolate; bracteoles as bracts but 


Sabiaceae (van Beusekom & van de Water) 


smaller, or bracteoles minute or sepal-like and then 
situated near calyx; pedicel up to 4 mm. Flowers 
green to yellow or white. Sepals sometimes 6 (see 
bracteoles), ± ovate or broad-ovate, 0.75- 1{- 1.25) 
by 0.5-0. 8(-l) mm, acute to obtuse, ± pubescent, 
ciliolate. Petals oblong, 2.5-3.5(-4) by 1-1.5 mm, 
obtuse, nerves up to 5, often dark-coloured and then 
conspicuous. Stamens (1-)1.25-1.5 mm; filament 
± flattened, (0.75-)l-1.25 mm long, 0.25-0.5 mm 
wide; anther globular to ellipsoid, 0.2-0.3 mm, in- 
flexed. Disk crown-shaped; ribs sometimes faint or 
absent. P/5/// 0.8-1.2 mm; style ± conical, 0.2-0.5 
mm, much shoner than the adjacent side(s) of the 
drupelet(s); ovary globular to subreniform, 0.5-0.6 
by 0.5-0.7 mm, glabrous. Drupelets obovoid or 
sometimes globular, ± compressed, 9-11 by 9-10 
mm, without persistent petals and stamens at the 
base; reticulate pattern often coarse and limited to 
the margin. Embryo with somewhat undulated or 
faintly folded cotyledons. 

Distr. Malesia: Sumatra (East Coast Res., In- 
dragiri, Lampongs), W. Java. In all c. 30 collections. 

Ecol. Forests, at (20-)200-1500 m. Ft. fr. 
Jan. -Dec. 

Vern. Java: areuj bebentjojan, a. kahawatang, 
a. katjapi, S. 

Notes. Sabia javanica strongly resembles S. 
pauciflora from the Philippines, the Moluccas, New 
Guinea, and the Solomon Islands. It can be 
distinguished from that species by its often more- 
flowered cymes, its shorter style, and some other 
slight differences. Since both species are geographi- 
cally separated, it was also possible to combine them 
into one species and give them the rank of subspecies. 
Although the differences are rather small, 1 believe 
that S. javanica and 5. pauciflora represent two dif- 
ferent, well-delimited, but very closely related spe- 
cies. Moreover, a reduction of both species to a single 
one would increase the variability of several taxo- 
nomic important characters, in consequence of 
which the delimitation with some other related 
species, like S. parvijlora and 5. racemosa, and 
possibly also S. limoniacea, would become less 
distinct. Finally, this might result into a far-going 
lumping and a reduction of all these species to, say, 
subspecies. Contrary to the situation in the extra- 
Malesian species .S. campanuiata Wall., however, in 
this case I believe that the differences between these 
taxa have reached a higher level already, resulting in 
the distinction of mutually closely related but ± 
well-delimited species, each with its own specific 
combination of characters. 

In vegetative characters and in drupelets S. 
javanica resembles .S. racemosa from Borneo. It can, 
however, easily be distinguished from that species by 
its morc-flowcred inflorescences and its floral 
characters, es{>ccially its petals. 

3. Sabia limoniacea Wall. [Cat. (1829) n. 1000, 
nom. nud.] ex Hook./. & Th. Fl. Ind. 1 (1855) 210; 
Walp. Ann. 4 (1857) 139; Benth. Fl. Hongk. (1861) 
70; Hook./. Fl. Brit. India 2 (1876) 3; Kurz, J. As. 
Soc. Beng. 45, ii (1876) 204, excl. syn. Sabia sp. 
Griffith (= S. parvijlora ssp. parviflora); For. Fl. 
Burma 1 (1877) 300 {'timonaceay, Forbes & 
Hemsley, J. Linn. Soc. Bot. 23 (1886) 144; King, J. 
As. Soc. Beng. 65, ii (1896) 454; Prain, Beng. PI. I 
(1903) 246; Brandis, Indian Trees (1906) 194; Dunn 
& Tutcher, Kew Bull. Add. Ser. 10 (1912) 68; 
Ridley, Fl. Mai. Pen. 1 (1922) 513; Merr. Lingnan 
Sc. J. 5 (1927) 19; Kanjilal c.s. Fl. Assam 1,2 (1936) 
326; Chen, Sargentia 3 (1943) 56, f. 7; Biswas, PI. 
Darj. Sikkim Himal. 1 (1966) 261; van de Water, 
Blumea 26 (1980) 44, / 6b, 8. - Androglossum 
reticulatum Champ, ex Benth. Hook. J. Bot. Kew 
Gard. Misc. 4 (1852) 42; Benth. Fl. Hongk. (1861) 
70; Chen, Sargentia 3 (1943) 58, non S. reticulata 
Elmer (1909). - Sabia celastrinea Muell. in Walp., 
Ann. 6 (1865) 1269. - Sabia malabarica Bedd. Ic. 
PI. Ind. Or. 1 (1874) 39. t. 177; HooK./ Fl. Brit. In- 
dia 2 (1876) 2; Brandis, Indian Trees (1906) 194; 
Gamble, Fl. Pres. Madras 1 (1918) 254; Chen, 
Sargentia 3 (1943) 48. - Fig. 2, 3. 

Evergreen woody climber, up to 10 m. Twigs 
glabrous or sometimes sparsely pubescent; flowering 
twigs up to 5 mm diam., glabrous to lax-pubescent. 
Buds broad-ovoid to ovoid, up to 2.5 mm, acute; 
scales (sub)glabrous, often ciliolate. Leaves oblong- 
ovate to lanceolate, 4-18 by 1.5-6.5(-8) cm, index 
2-4(-4.5), ± pergamentaceous-coriaceous, above 
and beneath glabrous or with some hairs especially 
on midrib; base acute to rounded, apex acute, 
sometimes obtuse, acuminate or not; nerves 5-9 
pairs, ± patent, sometimes somewhat ascending, 
curved to straight; petiole up to 2.5 cm, glabrous to 
lax-pubescent. Cymes either solitary, axillary, 
subtended by small and often herbaceous leaves, or 
when either the leaves are fallen or the cymes are 
subtended by bracts arranged in an up to 1 5 cm long, 
glabrous to ± lax-pubescent or tomentellous, race- 
mose to thyrsoid inflorescence, cymes up to 2 cm, 
l-4(-6)-nowered; pedicels up to 7 mm; bracts 
oblong, up to 4 mm, glabrous to pubescent, ciliolate; 
bracteoles ovate to oblong, up to 1 .75 mm, glabrous 
to pubescent, ciliolate, often situated near calyx. 
Flowers green to yellow or white. Sepals sometimes 
6 or 7 (see bracteoles), broad-ovate to elliptic, 
0.7- 1 .2(- 1 .5) by 0.6- I mm, acute to rounded, gla- 
brous to ± pubescent, ciliolate. Petals suborbicular 
to elliptic or ± obovate, 1.75-2.5 by 1.25-2 mm, 
obtuse to rounded, sometimes broad-acute, nerves 5, 
usually obscure. Stamens 1.5-2 mm; filament 
somewhat flaticncd, 1 .25- 1 .75 by 0.3-0.4 mm; an- 
ther ellipsoid, 0.25-0.35 mm, inflexed. Disk crown- 
shaped, thin; ribs often faint or absent. Pistil 


Flora Malesiana 

[ser. I, vol. lO^ 


Fig. 2. Sabia limoniacea Hook./. & Thoms. a. Habit, x2/3; b. ditto, with axillary cymes, x2/3; c. open 
flower, x4; d. petal and the opposed stamen, x 8; e. disk and pistil, x8 (a & c-e C.W. Wang 79409; b 

Wallich 1000). 


Sabiaceae (van Beusekom & van de Water) 


Fig. 3. Sabia limoniacea Hook./. & Thoms. a. fruit; 

a', embryo, both xl.5 (a Poilane 24769; a' 

POILANE 18918). 

0.7-1.2 mm; style conical to cylindrical, 0.2-0.6 
mm, much shorter than the adjacent side(s) of the 
drupelet(s); ovary globular to subreniform, 0.5-0.6 
by 0.5-0.8 mm, glabrous. Drupelets globular to 
obovoid, strongly compressed, 11-14 by 10-13 mm, 
red to blue or black when fresh, without persistent 
petals and stamens at the base; reticulate pattern 
usually faint or absent, sometimes more prominent 
at the margin. Embryo with somewhat undulated 

Distr. Continental SE. Asia (throughout India, 
Burma, Bangladesh, Thailand and Indochina to 
China); in Malesia: Malay Peninsula (incl. also P. 
Penang), Central Sumatra and Borneo (Sarawak), in 
all 7 collections. 

Ecol . Thickets and forest, 300-1200 m altitude. 
FL Sept. -Jan.,//-. Dec. -April. 

4. Sabia parvinora Wall, in Roxb., Fl. Ind. 2 (1824) 
310; G.Do.M, Gen. Hist. 2 (1832) 69; Walp. Rep. 1 
(1842) 557; Hook./ & Th. Fl. Ind. 1 (1855) 210; 
Walp. Ann. 4 (1857) 139; Hook./ Fl. Brit. India 2 
(1876) 2; Stapf, Trans. Linn. Soc. Lond. 4, 2 (1894) 
142; Bra.ndis, Indian Trees (1906) 194; Lecomte, Fl. 
Gen. I.-C. 2 (1908) 2, incl. var. harmandiana Le- 
comte, Bull. Soc. Bot. Fr. 54 (1907) 674; Kanjilal 
c.s. Fl. Assam 1, 2 (1936) 325; Chun, Sunyatsenia 4 
(1940) 242; Merr. Brittonia 4 (1941) 112; Chen, 
Sargentia 3 (1943) 64; Gagnep. & Vidal, Fl. Camb. 
Laos, Vietnam 1 (1960) 16; Biswas, PI. Darj. Sikkim 
Himal. 1 (1966) 261; Sen Gupta, Bull. Bot. Soc. 
Bcng.22, ii(1968) 196; Hara. Fl. E. Himal. 2 (1971) 
74; Sen Gupta, Rec. Bot. Surv. India 20, 2 (1973)65; 
Hah A & Williams, Enum. Fl. PI. Nepal 2(1970) 100; 
VAs de Water. Blumea 26 (1980) 48, f. 3c, 9. - 
\al)ia harmandiana Pierre, Fl. For. Coch. 5 (1897) 
pl 360B; Craib, Fl. Slam. Enum. I (1926) 340. - 
Sahia phitippinensis Robins. Bull. Torrcy Bot. Club 
' I9(J8) 70; .Merr. Enum. Philip. 2 (1923) 516; 

•.. Sargentia 3 (1943) 67. 

"■ a complete synonymy, see van de Water 


crgrccn climber or scandcnt shrub, up to 6 m. 
'.'."> glabrous to laxly pubescent; flowering (wigs 

up to 4 mm diam., glabrous to pubescent. Buds 
broad-ovoid to ovoid, up to 2 mm, acute; scales 
glabrous to short-pubescent, ciliolate. Leaves oblong 
to (sub)lanceolate, 3-12(-15) by 1-5 cm, index 2-4 
(-4.5), ± pergamentaceous, above glabrous to 
subglabrous or sometimes sparsely pubescent espe- 
cially when young, beneath glabrous to lax-pubes- 
cent especially on midrib; base acute to rounded, at- 
tenuate or not; apex acute, acuminate; nerves (5-) 
6-9(-10) pairs, patent, straight or sometimes ± 
curved; petiole up to 1.5 cm, glabrous to mainly 
above lax-pubescent. Cymes solilary, axillary, 1.5-8 
(-10) cm long, 4-25-flowered, sometimes widely 
spreading, lax, and with up to 35 or more flowers, 
glabrous to sparsely pubescent; pedicels up to 1 cm; 
bracts ovate to lanceolate, up to 2 mm or, when sub- 
tending a cyme up to 6 mm, subglabrous to pubes- 
cent, ciliolate; bracteoles as bracts. Flowers green to 
yellow or white. Sepals broad-ovate to ovate, 
0.7- 1 .5 by 0.5- 1 mm, acute to rounded, glabrous to 
pubescent, ciliolate. Petals elliptic-oblong to lanceo- 
late or sometimes oblong-ovate, 2-4(-4.5) by 
0.7-1.3 mm, acute to obtuse; nerves up to 7, dark- 
coloured or sometimes obscure. Stamens 1.2-2.25 
(-2.5) mm; filament flattened, 0.9-2(-2.25) by 
0.25-0.5 mm; anther ellipsoid to ovoid, 0.25-0.4 
mm, often ± inflexed. Disk crown-shaped, usually 
thin; lobes often distinct, relatively long and narrow, 
sometimes short or margin of disk irregular; ribs 
often faint or absent. Pistil l-2(-2.5) mm; style 
either absent or obscure, or conical, (0.75-)l-1.5 
(-1.75) mm, much shorter than the adjacent side(s) 
of the drupelet(s); ovary globular to subreniform, 
0.4-0.7 by 0.5-0.75 mm, glabrous. Drupelets glob- 
ular to somewhat obovoid, ± compressed, 7-9 by 
6-8 mm, green to red or blue when fresh, without 
persistent petals and stamens at the base; reticulate 
pattern rather fine, but often inconspicuous or ob- 
scure. Embryo with faintly wrinkled cotyledons. 

Distr. Widely ranging in SE. Asia from Nepal to 
China; in Malesia: N. Borneo (Sabah) and the Philip- 
pines (Luzon). 

KEY TO the subspecies 

I. Style normally developed, distinctly conical, 
(0.75-)l-1.5(-1.75) mm long 

a. ssp. parviflora 
1. Style usually absent or obscure, the upper part of 
the pistil carpel-like, sometimes normally devel- 
oped and then up to 0.75 mm 

b. ssp. philippinensis 

a. .w/7. parviflora - Sabia parviflora Wall. - Sabia 
harmandiana Pii.rri. . 

leaves oblong, somclimcs oblong-ovatc to (sub) 
lanceolate, 3~12(-I5) by 1-5 cm. C>mfi 2-8(-I0) 


Flora Malesiana 

[ser. I, vol. 104 

cm long, 7-25-flowered, sometimes widely spread- 
ing, lax, and with up to more than 35 flowers. Petals 
oblong to lanceolate, sometimes oblong-ovate, 
2.25-4(-4.5) by 0.7-1.25 mm. Style distinctly con- 
ical, (0.75-)l-1.5(-1.75) mm long. 

Distr. SE. Asia; in Malesia: Borneo (Sabah), 9 

Ecol. Roadsides, in thickets, and in forests, 
mainly 600-2000 m altitude. Fl. Jr. probably 
throughout the year. 

b. ssp. philippinensis (Robins.) van de Water, 
Blumea 26 (1980) 50. - Sabia philippinensis Robins. 
- Fig. 4. 

Leaves oblong or oblong-ovate to lanceolate, 
3-11 by 1-3.5 cm. Cymes 1.5-4.5 cm, 4-20-flow- 
ered. Petals elliptic-oblong to sublanceolate, 2-3.5 
by 1-1.25 mm. Style absent or obscure and often 
carpel-like, sometimes normal-developed and then 
up to 0.75 mm. Fruits not seen. 

Fig. 4. Sabia parviflora ssp. philippinensis (Robin- 
son) VAN DE Water, a. & b. disk and pistil showing 
the absence of a style; c. a feebly developed one; all 
X 12 (fl Ramos 26973; b Jacobs 7402; c Merrill 

Distr. Malesia: Philippines (Luzon: Benguet 
Prov.), 11 collections. 

Ecol. Forests, 71000-2100 m. Fl. mainly 
Febr. -April. 

Vern. Baybayok, kopdas, uakal, udok, Ig. 

Notes. Ssp. philippinensis can be distinguished 
rather easily from ssp. parviflora by the absence of 
a normally developed style. In all the specimens I 
have seen (except one) the upper parts of the two 
carpels of each flower are not connate with each 
other and differentiated into a style as usual, but re- 
main free and carpel-like, although the tip of each 
carpel is sometimes slightly stigmatic. Moreover, the 
margins of the upper part of a carpel are not fused, 
so that the upper half of each carpel remains open. 
Although this phenomenon is unique within the 
genus, I have reduced S. philippinensis to a subspe- 
cies of S. parviflora because it agrees very well with 
that species in all other main characters. 

Like in all Sabia species the leaves are dark above, 
paler beneath, but in the present one the contrast is 

especially conspicuous. In ssp. philippinensis the 
pale margins and undersides of the leaves provide a 
useful character to distinguish vegetative specimens 
from those of S. pauciflora, another Philippine 

5. Sabia pauciflora Blume, Mus. Bot. Lugd.-Bat. 1 
(1851) 370; MiQ. Fl. Ind. Bat. 1, 2 (1859) 619; Fl. 
Arch. Ind. (1870) 72; ibid. (1871) pi. 32; Chen, 
Sargentia 3 (1943) 61; van de Water, Blumea 26 
(1980) 51. - Sabia papuana Warb. in K.Sch. & 
Laut., Fl. Deut. Schutzgeb. Sudsee (1900) 425. - 
Sabia reticulata Elmer, Leafl. Philip. Bot. 2 (1909) 
579; Merr. Enum. Philip. Fl. PI. 2 (1923) 516; 
Chen. Sargentia 3 (1943) 62. 

Evergreen woody climber or scandent shrub, up to 
20 m. Twigs glabrous; flowering twigs up to 5 mm 
diam., glabrous or sparsely pubescent. Buds ovoid, 
up to 2.5 mm, acute; scales glabrous to pubescent, 
(sub)ciliolate. Leaver oblong to sublanceolate, 5-14 
(-18) by 2-6(-8) cm, index (2-)2.5-3.5(-4), above 
and beneath glabrous or with very few hairs on 
midrib, pergamentaceous; base acute to rounded, 
apex acute, acuminate; nerves (5-)6-8(-9) pairs, 
patent, straight to curved; petiole up to 2 cm, gla- 
brous to sparsely pubescent. Cymes either arranged 
in an axillary, up to 12 cm long, glabrous to sparsely 
pubescent, racemose to thyrsoid inflorescence, sub- 
tended by bracts, or solitary, axillary, often subtend- 
ed by small leaves, up to 3.5 cm, 1-4-flowered, gla- 
brous to sparsely pubescent; bracts oblong to 
lanceolate, up to 3.5 mm, subglabrous to somewhat 
pubescent, (sub)ciliolate; bracteoles as bracts but 
smaller, or minute, or sepal-like and then often 
situated near calyx; pedicels up to c. 1 cm. Flowers 
green to yellow or white. Sepals sometimes 6 (see 
bracteoles), ovate to broad-ovate, 0.75-1.25 by 
0.7-1 mm, acute to obtuse, glabrous to somewhat 
pubescent, (sub)ciliolate. Petals oblong, sometimes 
somewhat oblong-ovate, 2.5-4(-4.5) by (0.75-) 
1-1.3(-1.5) mm, (narrow-)obtuse, sometimes sub- 
ciliolate, nerves up to 5, sometimes dark-coloured 
and then conspicuous. Stamens (l-)1.25-1.75(-2) 
mm; filament flattened, (0.75-)l-1.5(-1.75) by 
0.25-0.5 mm; anther globular to ellipsoid, 0.2-0.3 
mm, inflexed. Disk crown-shaped; lobes often short 
or irregular; ribs sometimes faint or absent. Pistil 
1.3-1.7 mm; style conical, 0.6- 1 mm, much shorter 
than the adjacent side(s) of the drupelet(s); ovary 
globular to subreniform, 0.5-0.7 by 0.5-0.8 mm, 
glabrous. Drupelets ± globular, sometimes some- 
what obovoid, compressed, 7.5-11 by 8-10(-ll) 
mm, white to red or dark-blue when fresh, without 
persistent petals and stamens, reticulate pattern fine 
to rather coarse, sometimes indistinct, limited to the 
margin or not. Embryo with somewhat undulated or 
faintly folded cotyledons. 


Sabiaceae (van Beusekom & van de Water) 


Distr. Malesia: Moluccas (Buru, Halmaheira, 
Batjan), Philippines (Luzon, Negros, Mindanao), 
New Guinea; Solomon Islands. 

Ecol. Forests, from sea-level up to 2300 m. 
throughout the year. 

Uses. Fresh leaves eaten against wound fever in 
New Guinea. 

Vern. Philippines: bungoi, dadabu. Bag.; New 
Guinea: hambui, Poio, Enga lang., kubiakan, 
Hagen-Chimbu, Yoowi dial., mongotya ka. North- 
ern Prov., pehkuma, Mumuni, Orokaiva lang, pipi, 
E. Highlands, pukhabu, S. Highlands. 

Note. This species is closely related to S. javanica 
from Java and Sumatra, but can be distinguished 
from that species by its always few-flowered cymes, 
its longer style, and its often ± globular drupelets (S. 
javanica often obovoid). 

6. Sabia racemosa Chen, Sargentia 3 (1943) 36, f. 2; 
VAN DE Water, Blumea 26 (1980) 54. 

Evergreen woody climber or scandent shrub, up to 
6 m. Twigs glabrous; flowering twigs up to 4 mm 
diam., glabrous or somewhat short-pubescent. Buds 
ovoid, up to 1.5 mm acute; scales (sub)ciliolate or 
not. Leaves ob\ong or somewhat oblong-ovate, 6-25 
by 2-10 cm, index 2-3(-3.5), pergamentaceous, 
glabrous or with some hairs on midrib, rarely 
beneath all over sparsely short-pubescent; base acute 
to rounded, apex acute, acuminate; nerves 4-8 (or 9) 
pairs, ± patent, curved to straight; petiole up to 2.5 
cm, glabrous or with some very short hairs. Cymes 
arranged in an axillary, up to 8 cm long, glabrous to 
puberulous or short-tomentellous, racemose to thyr- 
soid inflorescence, subtended by bracts but often 
bracts fallen or sometimes leaf-like, cymes up to 1 
cm, I -4(-7)-nowered, glabrous to somewhat puber- 
ulous or short-tomentellous; bracts ovate to oblong, 
up to 3 mm, glabrous to somewhat pubescent, 
(sub)ciliolate; bracteoles as bracts but usually smal- 
ler, or minute and then often situated near calyx; 
pedicels up to 4 mm. Flowers (pale-)green to yellow. 
Sepals ± ovate to broad-ovate, 0.6-1.3 by 0.5-1 
mm, acute to obtuse, glabrous to somewhat pubes- 
cent, (sub)ciliolate. Petals elliptic-oblong to ovate- 
lanceolate, 3.5-6.5 by(1.25-)l.5-2.5 mm, acute to 
obtuse, or ± acuminate, or gradually narrowed, 
nerves up to 7, thin but distinct. Stamens 1.2-2.2 
mm; filament flattened, i -2 by 0.2-0.5 mm; anther 
globular to ellipsoid, 0.2-0.3 mm, inflexed. Disk 
crown-shaped; lobes sometimes very short or in- 
distinct; ribs sometimes faint or absent. Pistil i - 1.5 
mm; style ± conical, 0.5-0.9 mm, much shorter 
than the adjacent side(s) of the drupclct(s); ovary 
globular to subrcniform, 0.5-0.6 by 0.5-0.7 mm, 
glabrous. Drupe/e/5 obovoid, ± compressed, 10-12 
by (7- )8- 10 mm, while to pink or red when fresh, 
without persistent petals and stamens al the base; 

reticulate pattern faint to rather coarse, often limited 
to the margin. Embryo with somewhat to very 
wrinkled or folded cotyledons. 

Distr. Malesia: Borneo. 

Note. In vegetative characters and somewhat in 
the fruit this species resembles S. javanica. It differs, 
however, from that species in its inflorescence (few- 
flowered cymes) and in its floral characters, especial- 
ly the petals. 

Since the fruiting collections oi ssp. racemosa bear 
only immature or damaged fruit, the description of 
the drupelets has mainly been based on the fruit of 
ssp. kinabaluensis. 

The two subspecies can easily be distinguished 
from each other by the difference in the shape of 
their petals. Since they can be distinguished from 
each other only when flowers are available, the iden- 
tification of most of the vegetative and fruiting 
specimens has mainly been based on the locality from 
where they have been collected. 


1. Petals oblong-ovate to ovate-lanceolate, acute, 
somewhat acuminate or tapering to the apex 

a. ssp. racemosa 

1. Petals elliptic-oblong to oblong, acute to obtuse 

b. ssp. kinabaluensis 

a. ssp. racemosa - Sabia racemosa Chen. 

Sepals 0.6- 1.1 by 0.5- 1 mm. Petals oblong-ovate 
to ovate-lanceolate, (3.5-)4.5-6.5 by (1.25-)1.5- 
2.5 mm, acute, somewhat acuminate or tapering to 
the apex. Pistil 1-1.2 mm; style 0.5-0.7 mm long. 

Distr. Malesia: Borneo (Kalimantan), 7 collec- 

Ecol. Low altitudes, up to 100 m. Fl. fr. 
throughout the year. 

b. ssp. kinabaluensis van de Water, Blumea 26 
(1980) 55. 

Sepals 0.9-1.3 by 0.6-1 mm. Petals elliptic- 
oblong to oblong, 3.5-5 by 1.5-2.5 mm, acute to 
obtuse. Pistil 1.2-1.5 mm high; style 0.6-0.9 mm 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu), 
15 collections. 

Ecol. Forests, mainly al 800- 1 500 m altitude. Fl. 
fr. throughout the year. 

7. Sabia sumalruna Biumi., Mus. Bot. Lugd.-Bat. I 
(1851) 370; Miy. II. Ind. Bat. 1, 2 (1859) 619; Fl. 
Arch. Ind.(1870)72;;7>/ry. (187l)pl. 33; Kino, J. As. 
Soc. Beng. 65, ii (1896) 454; Ridley, Fl. Mal. Pen. 
1 (1922) 513; Chen, Sargentia 3 (1943) 39; van de 
Water. Blumea 26 (1980) 56. 

Evergreen woody climber, up to c. 3.5 m. Twigs 


Flora Malesiana 

[ser. I, vol. 10"^ 

glabrous; flowering twigs up to 4 mm diam., 
glabrous. Leaves elliptic to oblong, sometimes (sub) 
lanceolate, (5-)7- 15(- 18) by (1 .5-)2.5-7(- 10) cm, 
index 2-3(-4), pergamentaceous, above and be- 
neath glabrous; base acute, apex acuminate to sub- 
cuspidate; nerves 5-7 pairs, patent, curved to 
straight; petiole up to 2 cm, glabrous. Flowers yel- 
lowish-green to white, either solitary, sometimes 2 or 
3 together, axillary, or arranged in a thyrsoid, ax- 
illary, up to 6.5 cm long, glabrous inflorescence; 
pedicels up to 2.5 cm, glabrous, with few small 
budscales at the base when flowers solitary; bracts ± 
oblong-ovate, up to 1 .5 mm long, glabrous, ciliolate; 
bracteoles as bracts. Sepals broad-ovate to ovate, 
1.25-1.75(-2) by (0.75-)l-1.75 mm, acute to ob- 
tuse, glabrous, (sub)ciliolate or not. Petals oblong or 
ovate-lanceolate, c. 6- 10 by 1 .5-2.5 mm, sometimes 
the upper part somewhat channeled, tapering to the 
apex, acute to narrow-obtuse, nerves obscure. 
Stamens 2.5-1.5 mm; filament ± flattened, 3-7 by 
0.4-0.75 mm; anther ellipsoid, 0.5-0.7 mm, 
upright. Disk short-cylindrical, small, the upper part 
not enclosing the base of the ovary and without 
lobes; ribs ± prominent. Pistil 3.5-c. 7 mm; style 
narrow-conical, 3-6 mm, ± half as long as the adja- 
cent side(s) of the drupelet(s); ovary somewhat 

globular to subreniform, 0.5-0.8 by 0.7-1 mm, 
glabrous. Drupelets obovoid, somewhat compress- 
ed, 11-13 by 8-9 mm, white to blue when fresh, 
without persistent petals and stamens, reticulate pat- 
tern absent, often more or less rugged on the outside. 

Distr. Malesia: Sumatra (W. Coast Res., Palem- 
bang), 7 collections. 

Ecol. Forests, 60-1000 m altitude. Fl. May- 
Aug.,//-. July-Sept., Febr. 

Note. Only a few collections are available. For 
that reason no buds and embryos could be described, 
whereas the description of the flowers has partly 
been based on rather young ones. 


Sabia densiflora Miq. Sum. (1861) 203, 520 = 
Meliosma angulata Blume: K. & V. Bijdr. 9 (1903) 
131 = Meliosma simplicifolia (Roxb.) Walp. ssp. 
simplicifolia: van Beusekom, Blumea 19 (1971) 476; 
Fl. Males. 10" (1989) 698 (this issue). 

Sabia floribunda Miq. Sum. (1861) 203, 521 = 
Meliosma angulata Blume: K. & V. Bijdr. 9 (1903) 
131 = Meliosma simplicifolia (Roxb.) Walp. ssp. 
simplicifolia : /. c. 


Blume, Cat. (1823) 32; Rumphia 3 (1849) 196; Miq. Fl. Ind. Bat. 1 , 2 (1859) 612 
Benth. & Hook./. Gen. PI. 1 (1862) 414; Hook./. Fl. Brit. India 2 (1876) 3 
BoERL. Handl. Fl. Ned. Ind. 1 (1890) 290; Warb. in E. & P., Nat. Pfl. Fam 
3, 5 (1895) 371; van Beusekom, Blumea 19(1971)355. - Millingtonia Roxb 
[Hort. Beng. (1814) 3, nomen] PI. Corom. 3 (1820) 50, t. 254, non Linn./ 
(1781), nee Donn (1807). - Kingsboroughia Liebm. Vid. Medd. Nat. For 
Kjobenhavn 2 (1850) 67; Walp. Ann. 2 (1852) 834. - Fig. 5-8, 10, 12. 

For a complete synonymy, see van Beusekom (1971). 

Evergreen or sometimes deciduous shrubs or trees, up to 42 m, 1 m diam., 
sometimes buttressed. Twigs more or less lenticellate, often with conspicuous 
leaf-scars. Buds densely pubescent. Leaves simple or imparipinnate with 
(sub)opposite leaflets, ending in 3 or 1 leaflet(s), in the latter case its petiolule 
articulated with the rachis; leaves or leaflets entire or dentate, with or without 
hairy domatia beneath; rachis and petioles, usually also petiolules, with a usual- 
ly shallow and narrow, more or less conspicuous longitudinal groove above, 
usually with swollen base, articulately attached. Inflorescence terminal, some- 
times axillary, a pyramidal panicle, poor to usually profuse, up to 4 times 
ramified, with alternate, articulately attached, often lenticellate axes. Bracts 
small, those of lower order usually soon caducous; cataphylls often present. 
Bracteoles absent, but sometimes one (or two) bracteole-hke sepals present. 


Sabiaceae (van Beusekom & van de Water) 


Fig. 5. Flower of Meliosma. A. Semi-diagrammatical sketch of flower (subg. Meliosma) with opened outer 

petals, but stamens still in bud position. B. Semi-diagrammatical length section of bud (subg. Meliosma). C. 

Diagram {subg. Kingsboroughia and subg. Meliosma). Names of the flower parts: a. sepals; b. outer petals; 

c. inner petals; d. fertile stamens; e. staminodes; /. disk; g. style. 

lowered on the pedicel. Flowers numerous, sessile or short-pedicelled, small, 
bisexual. Sepals 5, by reduction sometimes 4, rarely 3, sometimes by addition 
of empty bracts seemingly more, up to c. 13, and together forming a kind of in- 
volucre, usually unequal and then mostly 3 about equal. Petals 5, episepalous, 
3 outer ones more or less unequal, alternisepalous, mostly suborbicular and con- 
vex, rarely the largest one much wider than long and more or less reniform, the 
smaller ones irregularly shaped; 2 inner ones equal, much smaller, reduced, op- 
posite the fertile stamens and more or less adherent to the base of the filaments, 
entire to bifid. Disk generally present, sometimes very reduced or absent, often 
irregularly shaped, as a rule with 5 more or less developed teeth, 4 of which 
paired, 1 unpaired, each pair opposite a fertile stamen. Stamens 5, epipetalous, 
2 fertile, filament short, strap-shaped, flat, incurved at the top, abruptly ter- 
minating in a wide, varyingly shaped cup which bears two globose to elliptic 
transversely dehiscent anther-cells which are ripe in bud, springing back elasti- 
cally when the flower opens; 3 staminodial, opposite the larger petals and more 
or less adherent to the base of these, deformed, broad, irregularly shaped, with 
1 or 2 holes near the top in which fit the anther-cells of the fertile stamens, often 
coherent and forming a cup over the pistil. Ovary globose to ovoid or conical, 
2-, very rarely 3-locular, apically contracted in a rather short, simple or 2- 
parlible, cylindric or subulate to conical, rarely minute style, with simple or 
somewhat bifid, minute stigma. Ovules 2 (or 1) in each cell, more or less 
superimposed, attached to the partition, hemi-anatropous. Fruit a drupe, 
subglobose to pyriform, small, glabrous, with one stone; rarely two ovules in- 
stead of one per ovary develop, resulting in a didymous fruit; mcsocarp pulpy, 
mostly thin; endocarp globose, pyriform, or semiglobose, 1-cellcd, stony to 
crustaceous, splitting in two valves, inside with a basilar rounded projection 
over which the seed is curved. Vascular bundle connecting pedicel and seed 
either running outside the endocarp wall (free in the pulpy mcsocarp or in a 
groove on the ventral endocarp wall), or through a canal inside the endocarp 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 6. Diagrammatical length sections of three types of fruit in Meliosma. A. Subg. Kingsboroughia sect. 
Hendersonia: vascular bundle running freely in the mesocarp. B. Subg. Kingsboroughia sect. Kings- 
boroughia: vascular bundle running in a groove of the endocarp, entering the wall through the ventral pore. 
C. Subg. Meliosma sect. Meliosma: similar to B, but the marginal canal lengthened through the endocarp. 

All x3. 

1989] Sabiaceae (van Beusekom & van de Water) 693 

wall. Seed sub- to semiglobose, more or less concave at the ventral side, with 
membranous testa, without endosperm. Embryo with rather long, 2-3 times 
folded radicle and more or less folded cotyledons. 

Distr. About 20-25 species, 15 of which in SE. Asia, and not more than c. 10 in Central and South 
America. In Malesia: 8 species. 

The New World species belong to Meliosma subg. Meliosma sect. Lorenzanea, a section restricted to the 
New World; besides, there is one species oi subg. Kingsboroughia which is widely spread in China but also 
occurs in Mexico (A/, alba Walp.). 

Correctly named fossils from the Tertiary are found widely distributed on the northern hemisphere, in 
Europe, Asia, and North America; see van Beusekom, I.e. 384-424, fig. 16-18 (maps). The oldest known 
fossils, of both subgenera, date from the Eocene. All localities lie south of the 60° parallel of latitude and 
almost all beyond the present range of the genus. It is remarkable that still in the Pliocene the genus occurred 
in Europe, S. Russia, but no longer in North America. Only in southern Japan Pliocene fossils and recent 
species are found together. 

Ecol. In primary and secondary forests, especially on hills and mountains up to c. 3300 m, but also in 
lowlands. All or almost all species prefer everwet to moist, tropical to subtropical conditions. Some are hardy 
in mild temperate climates; these are deciduous and grow flush-wise. 

Morph. Trees, mostly small, sometimes shrubs, rarely mentioned to be subscandent, but A/, pinnata ssp. 
ferruginea and ssp. macrophylla are recorded to reach 42 m height and M. lanceolata to reach 30 m by 1 m 

The margin of leaf or leaflet may be entire or dentate and is often variable. In saplings, watershoots and 
seedlings the margin is mostly dentate. In species with pinnate leaves the size of the leaflets mostly increases 
apically and their greatest width tends to shift towards the upper half. The leaves, when pinnate, have I or 
3 top-leaflets; in the first case the petiolule of the top-leaflet has an articulation with the rachis. 

The inflorescence consists of a racemosely arranged, rich-fiowered panicle. 

Van Beusekom {I.e. 361-364, fig. 2 & 3) amply discussed the peculiar fiower structure. Although Baillon 
assumed the fiower to be basically 3-merous, he agrees with the majority of authors that it is 5-merous. The 
3 outer petals are differently shaped from the 2 inner ones; the latter may be of the lanceolate or bifid type, 
and taxonomically their shape is important. 

The structure of endocarp and seed(/.c. 364-369, fig. 4)isof great importance. The ovary contains 4 ovules 
but only one develops into a seed (exceptionally 2, resulting in an anomalous didymous fruit). The fruit is 
a drupe with rather thin, pulpy mesocarp and a stony to crustaceous endocarp, more or less globular to 
pyriform, smooth or often with a reticulate surface. When dehiscent, it splits into two valves, the plane of 
dehiscence usually marked by a ± prominent keel running all around the endocarp. At the ventral side there 
is a usually narrow pore through which the seed is connected with the vascular bundle towards the pedicel. 
There are two main types: 1) endocarps which only enclose the seed, whereas the vascular bundle connecting 
pedicel and seed is running outside the endocarp wall; 2) endocarps which enclose both seed and vascular bun- 
dle, the latter being situated in a marginal canal inside. 

Taxon . The subdivision of the genus Meliosma is as follows: 

1. Leaves simple or pinnate; when pinnate rachis terminating in 3 leaflets (anomalously 2 or 1). Sepals mostly 
5. Outer petals narrowly imbricate, subrotund to broad-elliptic, all regularly shaped. Vascular bundle con- 
necting pedicel and seed situated in a long or short marginal canal inside the endocarp. About 12 species 

in SE. Asia. Spp. 1-7 Subg. Meliosmii 

2. Leaves simple or pinnate. Ovary glabrous or pubescent. Endocarp wall relatively thin, not drawn out 
around the ventral perforation; endocarp mostly (sub)globosc, sometimes semiglobose, or ellipsoid to 

obovoid. About 12 spp. in SE. Asia. Spp. 1-7 Sect. Meliosma 

3. Leaves simple. Spp. I A 2 Subsecl. Simplices (Warb.) Beus. 

4. Deciduous shrubs or small trees. Nerves all or almost all straight or almost straight, ("oniincnial Asia 

Ser. Krclinervia Beus. 
4. Evergreen shrubs or trees. Nerves all or almost all distinctly ascending. Spp. I A 2 

Ser. Curvinervia Beus. 

3. Leaves pinnate. Spp. 3-7 Subsect Pinnalae (Warb.) Beus. 

2. Leaves simple. Ovary always glabrous. Endocarp wall relatively thick, more or less drawn out around the 


Flora Malesiana [ser. I, vol. 10^ 

ventral perforation which often gives the mostly (sub)globose endocarp a somewhat pyriform shape. 

About 10 species in Central and tropical South America Sect. Lorenzanea (Liebm.) Beus. 

1 . Leaves pinnate, petiolule of terminal leaflet articulate with the rachis. Sepals mostly 4. Outer petals widely 
imbricate, the largest one widely reniform, much wider than long, the smaller ones of irregular shape ± 
not wider than long. Vascular bundle connecting pedicel and seed situated outside the endocarp, either run- 
ning in a groove at the ventral side or freely in the pulpy mesocarp 

Subg. Kingsboroughia (Liebm.) Beus. 
5. Deciduous trees. Vascular bundle connecting pedicel and seed running in a ventral groove of the endocarp 

wall. Two species. SE. Asia and S. Mexico Sect. Kingsboroughia 

5. Evergreen trees. Vascular bundle connecting pedicel and seed running freely in the mesocarp. One species. 
Malay Peninsula, N. Borneo. Sp. 8 Sect. Hendersonia Beus. 


1 . Leaves simple. Subg. Meliosma sect. Meliosma subsect. Simplices. 

2. Petioles 1/5-1/3 the length of the lamina. Panicles mostly axillary or sometimes ramiflorous, sometimes 
terminal. Inner petals bifid or entire. Endocarps ellipsoid to obovoid, with reticulate to smooth surface, 
rarely (sub)globose and then smooth, 6-14 cm diam 1. M. lepidota 

2. Petioles 1/20-1/5 the length of the lamina. Panicles always terminal. Inner petals always bifid. Endo- 
carps always (sub)globose, always with reticulate surface, 3.5-8 mm diam 2. M. simplicifolia 

1 . Leaves pinnate. Subg. Meliosma and subg. Kingsboroughia. 

3. Leaf-rachis terminating in 3 (sometimes 2, rarely 1) leaflets. Outer petals widely ovate to orbicular, entire. 
Endocarps inside with a marginal canal in which runs the vascular bundle connecting pedicels and seed. 
Sect. Meliosma subsect. Pinnatae. 

4. Leaves 2-5(-6)-jugate; leaflets usually glabrous, sometimes (in Bornean specimens) pubescent beneath, 
above always with a prominent midrib. Inner petals (1.2-)1.5-2(-3) mm, entire to retuse or slightly 

bifid at apex. Ovary glabrous. Endocarps 0.7-2 cm diam 3. M. sumatrana 

4. Leaves 2-23-jugate; leaflets glabrous or pubescent, midrib usually flat to sulcate above. Inner petals 
(0.3-)0.5-l(-1.5) mm, always distinctly and rather deeply bifid. Ovary glabrous or pubescent. En- 
docarps 0.2-1 cm diam. 
5. Leaves (3-)6-18(-23)-jugate, with (10-)20-100 cm long rachis. Leaflets only very rarely with slight 
pubescence on midrib and nerves above. Panicles large and lax, 0.5-1.5 m, pendulous, usually suddenly 
bent down at the base, with up to 90 cm long primary side-axes which are never subtended by (small) 
6. Leaflets in middle and upper part of the leaf elliptic to lanceolate, very rarely a few linear-lanceolate, 
index (0.5-)2-5(-7). Petiolules 2-30 mm 4. M. lanceolata 

6. Leaflets in middle and upper part of the leaf linear-lanceolate, index 5-10. Lateral petiolules absent 
or very short, to c. 1 mm 5. M. hirsuta 

5. Leaves 2-7(-9)-jugate, rachis up to c. 40(-60) cm. Leaflets usually more or less pubescent on midrib 
and nerves above, sometimes glabrous. Panicles lax to dense but not very large, 10-50(-70) cm, usually 
erect, sometimes ± pendulous, but almost never suddenly bent dov/n at the base, with up to 35(-60) 
cm long side-axes which may be subtended by decrescent leaves. 

7. Sepals pubescent. Outer petals glabrous. Leaves 2-3(-4)-jugate. Endocarps 7-8 mm diam. Small 
trees 7. M. sarawakensis 

7. Sepals glabrous (rarely with a few hairs), sometimes pubescent but then also outer petals pubescent. 
Leaves 2-7(-9)-jugate. Endocarps 3-9(-10) mm diam. Small to large trees. 

8. Plants from Malesia except Sumatra and Java 6. M. pinnata 

8. Plants from Sumatra and Java. 
9. Leaflets pubescent at least on midrib and nerves, usually entire. Panicles often with decrescent 

leaves. Lowland and mountains up to 2500 m 6. M. pinnata 

9. Leaflets (sub)glabrous, usually (obscurely) dentate. Panicles without decrescent leaves. Mountains, 

1300-2900 m 4. M. lanceolata/. nervosa 

3. Leaf-rachis terminating in 1 (or 2) leaflets of which the short petiolule is always well markedly articulated 
with the rachis. Outer petals mostly widely reniform, or of irregular shape, mostly wider than long, with 
± irregular margin and often emarginate. Endocarps inside without a marginal canal in which runs the 
vascular bundle, large, wider than 1 cm. Inner petals hardly or not incised at apex. Subg. Kingsboroughia 
sect. Hendersonia 8. M. rufo-pilosa 


Sabiaceae (van Beusekom & van de Water) 


1. Meliosma lepidota Blume, Rumphia 3 (1849) 199; 
Walp. Ann. 2 (1852) 224; MiQ. Fl. Ind. Bat. 1, 2 
(1859) 614; Sum. (1861) 203; van Beusekom, Blumea 
19(1971)451,/. 25. - Fig. 7. 

For further synonyms, see under the subspecies; 
for a complete synonymy, see van Beusekom (1971). 

Evergreen shrub or tree, up to c. 15(-22) m. Flow- 
ering twigs pubescent when young, glabrescent. 
Leaves elliptic or obovate to lanceolate, 2-32 by 
0.7-12(-18)cm, inde.\(1.2-)1.5-3(-4), atthebase 
acute, at apex acute to caudate, rarely obtuse, usual- 
ly entire, sometimes remotely spinously dentate to- 
wards the apex, beneath sometimes pubescent on 
midrib and nerves, without domatia; nerves 7-15 
pairs, usually strongly ascending, petioles usually 
rather long, 1-10 cm, 1/5-1/3 as long as the blade. 
Panicles usually axillary and erect, widely to usually 
narrowly pyramidal, 3-30(-200) cm, usually dense- 
ly pubescent, bearing numerous solitary to crowded 
flowers which are sometimes spicately arranged; 
side-axes usually many, usually short, up to c. 15 
(-40) cm, sometimes subtended by normal to small 
leaves; bracts ovate to linear-lanceolate, up to c. 
2(-6) mm, usually densely pubescent. Pedicels ab- 
sent or present, up to c. 3(-5) mm. Mature buds 
1.5-3 mm diam. Sepals (4) 5, (round-)ovate, sub- 
equal, 1 -2 mm, or the outer 1 or 2 smaller, often one 
lowered on the pedicel, all entire, ciliolate. Outer 
petals glabrous. Inner petals ± lanceolate and entire, 
or bifid, (0.6-)0.8-2.5 mm, glabrous or somewhat 
ciliolate at margin or tip, when bifid never with a cen- 
tral lobule. Filaments 0.7-1.5 mm. Ovary 0.5-1 
mm, very exceptionally pubescent. Fruit (sub)glo- 
bose, sometimes elliptic, when ripe 5-10 mm diam.; 
endocarp globose to ellipsoid, 6-8(-9) mm diam., 
usually with a slightly elevated rather fine reticulum; 
median keel distinct, more or less prominent; ventral 
pore whether or not sunken but never spouted. 

Distr. SE. & E. Asia; in Malesia (with 4 
subspecies): Sumatra, Malay Peninsula, W. Java, N. 
Borneo (Sabah), and the Philippines (Luzon, Min- 

Ecol . In evergreen forests under tropical or sub- 
tropical conditions, at medium to high altitudes; for 
details, see under the subspecies. 

Notes . Meliosma lepidota displays a rather wide 
variation, especially in the ramification of its pan- 
icles which covers almost the whole range of possibil- 
ities found throughout Meliosma. 

Within M. lepidota seven subspecies are recog- 
nized, four of which in Malesia. The differences be- 
tween them arc on the same level as in other sub- 
species in Meliosma. Transitional forms between 
these subspecies, however, occur in only a few cases, 
which is logical since there is perfect geographical 
isolation between most of them. Sec further the notes 
under the subspecies. 

key to the subspecies 

1. Inner petals distinctly bifid. 
2. Leaves 1 .5-2(-2.5) times as long as wide; petiole 
1/4-2/3 as long as the blade. Panicles 5-15 cm. 
Mature buds 2-2.5 mm diam. Endocarps ellip- 
soid to obovoid d. ssp. kinabaluensis 

2. Leaves (1.6-)2-3 times as long as wide; petiole 
(l/6-)l/5-l/3 (-1/2) as long as the blade. 
Panicles 3-30 cm. Mature buds 1 .5-2(-2.2) mm 
diam. Endocarps long- to short -ellipsoid (always 
distinctly higher than wide) ... .a. ssp. lepidota 

1. Inner petals entire, usually lanceolate. 

3. Inner petals 2.5 mm. Panicles distinctly axillary 
or ramiflorous. Mature buds 2.5-3 mm diam. 

b. ssp. dolichomischa 
3. Inner petals 1-1.5 mm. Panicles terminal or 
crowded at the end of the twigs, rarely distinctly 
axillary. Mature buds 2-2.5 mm diam. 

c. ssp. vulcanica 

a. ssp. lepidota. - Meliosma lepidota Blume, Rum- 
phia 3 (1849) 199; Walp. Ann. 2 (1852) 224; MiQ. Fl. 
Ind. Bat. 1, 2 (1859) 614; Sum. (1861) 203; Illustr. 
(1871) 73. - Meliosma pedicellata K. & V. Bijdr. 9 
(1903) 134; Koord. Exk. Fl. Java 2 (1912) 545; Atlas 
2 (1914) t. 379; Backer & Bakh./. Fl. Java 2 (1965) 

Leaves oblong, sometimes somewhat ovate- 
oblong, rarely elliptic, 5-26 by (1.5-)2-12 cm, en- 
tire, base acute, apex acute to caudate, glabrous 
when mature; nerves 8-12(-14) pairs; petiole 1.5-6 
cm. Panicles axiWary, rarely terminal or ramiflorous, 
often several together near the end of a branch, 3-30 
cm, rather poor and lax, ramified up to the 2nd 
order; primary (essentially secondary!) side-axes 
short, up to c. 6(-10) cm. Mature buds 1.5-2 mm 
diam. Inner petals about halfway bifid, 0.7-1 mm; 
lobes rather narrow. Endocarp obovoid to ellipsoid, 
(8-)9-14 mm long, 5.5-8 mm diam., with or 
without rather wide and feeble reticulum; median 
keel distinct, more or less prominent, blunt to rather 
sharp, at one or both ends running out into a ventral, 
often somewhat beak-like processus; ventral pore 
rather wide, somewhat sunken. 

Distr. Malesia: Sumatra (not uncommon in 
Aceh, Tapanuli, and West Coast), W. Java. 

Ecol . Primary montane rain-forest; 900-26(X) m 
altitude in Sumatra, 1050-1600 m in Java. 

Field notes . Outer bark dark brown, finely cor- 
ky, 0.5 mm; inner bark turning rcdbrown, 0.5 cm; 
wood ochre with reddish stripes. 

Vern . Sumatra: antuang, hontuang, Batak lang., 
Toba, kalompang bag^h, CJn. Talamau. 

Note. .Ssp. lepidota is similar and probably most 
closely related to the adjacent s.<!p. dolichomi.\cha 
and ,v.v/>. kinabaluensis. However, ssp. lepidota also 


Flora Malesiana 

[ser. I, vol. 104 

Fig. 7. Meliosma lepidota Blume ssp. dolichomischa (Vidal) Beus. a. Flowering twig; xO.5; b. half-opened 

flower, x4.5; c. outer petal with adhering staminode; d. stamen with adhering inner petal, adaxial view; e. 

stamen, abaxial view;/, pistil with surrounding disk; g. ovary, length section, all x9; h. fruit, x 1.5 {a-g 

Henderson SF 23488; h Henderson SF 23492). 

shows a close resemblance to certain forms of ssp. 
longipes (Merr.) Beus. from Vietnam, from which 
it can sometimes only be distinguished by the shape 
of the endocarp. 

b. ssp. dolichomischa (Vidal) Beus. Blumea 19 
(1971) 458, f. 25. - Meliosma dolichomischa Vidal, 
Not. Syst. 16 (1960) 304. - Meliosma monophylla 
Ridley, J. Str. Br. Roy. As. Soc. n. 54 (1910) 40, 
nom. illeg.,non Merr. (1909); Fl. Mai. Pen. 1 (1922) 

514; Vidal, Not. Syst. 16 (1960) 306. - Fig. 7. 

Leaves elliptic to oblong, 4-22 by 2-10 cm, en- 
tire, base mostly attenuate, apex usually cuspidate, 
glabrous or subglabrous, nerves 7-13 pairs, petiole 
(l-)3-10 cm. Panicles axillary or ramiflorous, 
solitary or a few together, 6-25 cm, rather poor and 
lax, ramified up to the 2nd or 3rd order; primary (es- 
sentially secondary!) side-axes up to c. 10 cm. 
Mature buds 2.5-3 mm diam. Inner petals lanceo- 
late, c. 2.5 mm, entire, hooding over the stamens, 


Sabiaceae (van Beusekom & van de Water) 



Fig. 8. Various types of inner petals in Meliosma simplicifolia Walp. a. ssp. pungens (Walp.) Beus., b. ssp. 
rigida (Sieb. & Zucc.) Reus., c. ssp. fruticosa ^lume) Beus., d. ssp. simplicifolia; all x 18. 

glabrous. Endocarp as in ssp. lepidota. 

Distr. Malesia: Malay Peninsula (Pahang: 
Eraser's Hill, Cameron Highlands; Perak: Hermi- 
tage Hill, once). 

Ecol. Primary montane rain-forest, c. 1200- 
1500 m altitude. 

Field notes. Bark thick, red. Wood first white 
when cut, darkening to orange-brown. Leaves 
glaucous below. 

c. ssp. vulcanica (Merr.) Beus. Blumea 19 (1971) 
460. - Meliosma vulcanica Merr. Philip. J. Sc. 11 
(1916) Hot. 15; Enum. Philip. Fl. PI. 2 (1923) 518. - 
Machilus nervosa Merr. Philip. J. Sc. 4 (1909) Bot. 
262; Enum. Philip. Fl. PI. 2 (1923) 189; Salvore & 
Lagrimas, Philip. J. For. 4 (1941) 309; c/. Kosterm. 
Reinwardtia 5 (1960) 377; Bibl. Laur. 1 (1964) 919. 
- .Meliosma boniocensis Merr. Philip. J. Sc. 20 
(1922) 403; Enum. Philip. Fl. PI. 2 (1923) 517; 
Kosterm. Reinwardtia 5 (1960) 377. 

Leaves obovate-oblong or oblong, 5-16 by 2-6 
cm, base acute, apex acute to acuminate or 
sometimes rounded, glabrous or subglabrous; nerves 
8-11 pairs; petiole 2-4 cm. Panicles terminal, 
sometimes axillary, 3-20 cm, rather profuse to poor, 
ramified up to the 3rd or 4th order; primary side-axes 
(mostly essentially primary!) up to c. 15 cm, usually 
subtended by normal to reduced leaves. Mature buds 
2-2.5 mm diam. Inner petals lanceolate, 1-1.5 mm, 
entire, sometimes frayed at the tip. Endocarp subglo- 
bosc, rather oblique, 6-7 mm diam., apart from a 
few ribs smooth, median keel distinct, rather promi- 
nent, at one end running out into a minute ventral 
processus; ventral pore somewhat sunken. 

Distr. Malesia: Philippines (Luzon, Mindoro). 

Ecol. Primary rain-forest, low altitude up to c. 
2000 m. 

Note . Ssp. vulcanica is the only subspecies of M. 
lepidota in which normal terminal panicles have been 
found. In general habit it is more similar to certain 
forms of ssp. longipes from Vietnam than to .ssp. 
squamulata (Han* e) Bius. from Taiwan or to ssp. 
kinabatuensis from Borneo, to which i( is obviously 
less closely related. 

d. ssp. kinabaluensis Beus. Blumea 19 (1971) 455. - 
Meliosma pedicellata {nan K. & V.) Merr. & Perry, 
J. Arn. Arb. 20 (1939) 356. 

leaver elliptic, rarely oblong, 3-15 by 1.5-9 cm, 
usually entire, base acute to rounded and somewhat 
attenuate, cuspidate, above glabrous or ± pubescent 
on the midrib, subglabrous beneath, usually with a 
white waxy layer beneath, which gives a glaucous ap- 
pearance; nerves 8-14 pairs; petiole 1-7.5 cm. 
Panicles terminal or axillary, solitary or a few togeth- 
er, 5-15 cm, rather poor and lax, ramified up to the 
2nd (3rd) order; primary (essentially secondary!) 
side-axes up to c. 8 cm. Mature buds 2-2.5 mm 
diam. Inner petals halfway or somewhat less bifid, 
1-1.2 mm; lobes rather narrow. Endocarp ± 
obovoid, c. 8 mm long, c. 6 mm diam., with rather 
wide and feeble reticulum; median keel only slightly 
elevated, blunt, at one end running out into a minute 
ventral processus; ventral pore wide, not sunken. 

Distr. Malesia: Borneo (Mt Kinabalu). 

Ecol. Montane forest, 1700-2700 m altitude. 

Field notes. The lower surface of the leaves is 
often said to be white to light grey; in herbarium 
specimens indeed a whitish waxy layer can be ob- 
served often. The general colour of the leaves is re- 
ported to be glaucous. 

Note. Ssp. kinabaluensis has a very low degree of 
variability, a characteristic which is also found in 
some other subspecies of M. lepidota. It is most 
similar to ssp. dolichomischa from the Malay Penin- 
sula and to ssp. lepidota from Sumatra, with which 
it shares, amongst others, the more or less ellipsoid 
endocarp; all other subspecies have (sub)globosc en- 

2. Meliosma simplicifolia (Roxb.) Waip. Rep. 1 
(1842) 103; HA.ssK.Cat. Hon. Bog. (1844) 226; Miy. 
Fl. Ind. Bat. 1, 2 (1859) 613; Sum. (1861) 203; van 
BhUSEKOM, Blumea 19(1971)462, f. 28. Milling- 
tonia simplicifolia Roxb. [Horl. Bcng. (1814) 3, 
nomen] PI. Corom. 3 (1820) 50, t. 254. FIr. 8. 10. 

For further synonyms, see under the subspecies; 
for a complete synonymy, see van Beusekum ( 1 97 1 ). 

Evergreen shrub or tree, up to 20 m. Leaves elliptic 


Flora Malesiana 

[ser. I, vol. 104 

or obovate to lanceolate. 3-50 by 1-18 cm, base 
cuneate, apex acute to acuminate, rarely caudate or 
rounded, entire to spinously dentate, sometimes with 
hairy domatia; nerves 7-25 pairs, ± ascending, 
sometimes looped; petiole 0.5-6(-7) cm, 1/20-1/3 
as long as the blade. Panicles terminal, very rarely 
axillary, erect, lax to rather dense, widely to narrow- 
ly pyramidal, (4-)10-50(-60) cm, usually profusely 
branched up to the 2nd-4th order, bearing numer- 
ous solitary to crowded or glomerulate flowers which 
are usually spicately arranged; primary side-axes 
usually many, up to c. 25 cm, often subtended by 
leaves; bracts ovate to linear-lanceolate, up to c. 8 
mm. Pedicels sometimes present, up to c. 3 mm. 
Mature buds (1-) 1.5-3 mm diam. Sepals (4) 5, 
sometimes by addition of empty bracts seemingly 
more, up to 11(-13), (round-)ovate, equal or usually 
more or less unequal, the inner ones 0.7-2 mm, the 
outer one(s) smaller, often minute. Inner petals more 
or less deeply bifid, 0.5-1.5 mm, with glabrous, 
sometimes fimbriate or ciliolate lobes, never with a 
central lobule. Filaments 0.5-1.5 mm. Ovary 
0.5-0.7(-l) mm. Mature fruit (sub)globose, 4-10 
mm diam.; endocarp globose to subglobose, often 
depressed or oblique, 3-9 mm diam., with very 
vague to very strong and prominent reticulum; me- 
dian keel more or less prominent; ventral pore 
somewhat sunken to somewhat spouted. 

Distr. Continental SE. Asia (from Ceylon to 
China, Taiwan and S. Japan); in Malesia: Sumatra, 
Malay Peninsula, Borneo, Java, and Lesser Sunda 
Islands. Fig. 9. 

Ecol. Subtropical to tropical forests, under 
various conditions, usually in mountains up to c. 
3000 m, but also at sea-level. For details see under the 

Note. Meliosma simplicifolia is a very variable 
species, covering an enormous area in which it is 
adapted to many different habitats. It can be divided 
into eight well-marked subspecies, five of which cen- 
tre in SW. Yunnan, and diverge over different parts 
of the area. 


1. Sepals (4-)5. 

2. Panicles branched up to the 2nd (3rd) order, 
nearly always (very) densely tomentose; primary 
side-axes rarely subtended by leaves. Leaves 
sparsely to densely but always distinctly pubes- 
cent to tomentose, at least on midrib and nerves; 
without domatia. Style c. 1.5-2 times as long as 
the ovary b. ssp. rigida 

2. Panicles branched up to the (2nd) 3rd or 4th 
order, sparsely pubescent to moderately tomen- 
tose; lower primary side-axes often subtended by 
normal to small or reduced leaves. Leaves gla- 

brous to densely pubescent, rarely tomentose, 

with or without domatia. Style about as long as 

the ovary or shorter. 

3. Leaves with or without domatia; midrib on the 

upper side of the full-grown leaf glabrous or 

nearly so, more or less prominent, rarely flat. 

Inner petals with entire lobes, which are 

sometimes slightly fimbriate or ciliolate at the 

very tips. Endocarps 3.5-5(-7) mm diam. 

a. ssp. simplicifolia 
3. Leaves with or without domatia; midrib on the 
upper side of the full-grown leaf more or less 
but distinctly pubescent, ± impressed to flat. 
Inner petals usually with fimbriate, rarely entire 
lobes which are rarely minutely ciliolate at the 
very tips. Endocarps (4.5-)5.5-8 mm diam. 

c. ssp. fruticosa 
1. Sepals (8-)9-ll(-13). Leaves usually with do- 
matia. Endocarps 3.5-5.5 mm diam. 

d. ssp. pungens 

a. ssp. simplicifolia. - Millingtonia simplicifolia 
RoxB. [Hort. Beng. (1814) 3, nomen] PI. Corom. 3 
(1820) 50, t. 254; Fl. Ind. 1 (1820) 103; Nees, Flora 

8 (1825) 106; Griff. Not. PI. As. (1854) 162; Ic. PI. 
As. (1854) t. 442; Anon. Ic. Roxb. 4 (1970) 40, t. 20; 
VAN Beusekom, Blumea 19 (1971) 476. - Meliosma 
simplicifolia Walp. Rep. 1 (1842) 103; Hassk. Cat. 
Hort. Bog. (1844) 226; Thw. Enum. PI. Zeyl. (1858) 
59; MiQ. Fl. Ind. Bat. 1, 2 (1859) 613; Sum. (1861) 
203; Illustr. (1871) 73; Bedd. Fl. Sylv. 3 (1871) 77; 
Brandis, For. Fl. (1874) 116; Hook./. Fl. Brit. India 
2 (1876) 5; KuRZ, J. As. Soc. Beng. 45, ii (1876) 204; 
Fl. Burma 1 (1877) 301; Trim. Fl. Ceyl. I (1893) 315; 
Prain, Bengal PI. 1 (1903) 246; Brandis, Indian 
Trees (1906) 194; Merr. Contr. Arn. Arb. 8 (1934) 
95; Brittonia 4 (1941) 110; Vidal, Not. Syst. 16 
(1960) 307. - Meliosma angulata Blume, Rumphia 3 
(1849) 197; Walp. Ann. 2 (1852) 224; K. & V. Bijdr. 

9 (1903) 131; Koord. Exk. Fl. Java 2 (1912) 545; 
Atlas 2 (1914) t. 378; Baker/, in Rendle, J. Bot. 62 
(1924) Suppl. 30; Vidal, Not. Syst. 16 (1960) 304. - 
Sabia densiflora Miq. Sum. (1861) 203, 520. - Sabia 
floribunda Miq. I.e. 203, 521; Kurz, J. As. Soc. 
Beng. 39, ii (1870) 74. - Fig. 8d. 

Leaves obovate-oblong to -lanceolate, up to c. 50 
by 18 cm, base cuneate, apex acute to short-cuspi- 
date, beneath often with domatia; nerves 8-23 pairs. 
Panicles rather lax, 10-45 cm, branched up to the 
3rd or 4th order; axes sparsely to densely pubescent 
but never tomentose, the lower primary ones sub- 
tended by leaves. Flowers more or less crowded to 
solitary, (sub)sessile; mature buds 1.5-2 mm diam. 
Sepals 5 (4). Inner petals 0.6-0.8 mm, usually over 
halfway bifid, lobes more or less divergent, narrow, 
glabrous, sometimes slightly fimbriate or ciliolate at 
the very tips. Style about as long as ovary or shorter. 


Sabiaceae (van Beusekom & van de Water) 


M. simplici folia 

^^m ssp.fordii 
• •M ssp. rigida 
_^_ ssp. thomsonii 
^^= ssp. laui 

ssp. frulicosa 

ssp. pungens 

._- ssp.yunnanensis 

ssp. simplicifolia 

Fig. 9. Generalized areas of the subspecies of Meliosma simplicifolia Walp. 

Endocarps subglobose, usually rather oblique, near- 
ly triangular at ventral view, 3.5-5(-7) mm diam., 
with more or less prominent, rather coarse reticu- 
lum; median keel usually very prominent, at one end 
sometimes running out into a minute ventral pro- 
cessus; ventral pore somewhat or not sunken, not 

Distr. Widely distributed in continental SE. 
Asia; in Matesia: northern half of Sumatra, W. Java 
(not found since Blume's time). Fig. 9. 

Ecol. Primary and secondary evergreen forest, 
from sea-level up to c. 1200<- 1500) m altitude. It is 
often reported to occur along watercourses. 

Vern. Sumatra: medanf; sungu, M, simulingga, 
sumpa mana belawah, Karo, kayu gadis. West 

Note. A rather uniform, well recognizable sub- 
species all over its area. 

b. ssp. rigida (Sieb.&Zucc.)Beus. Blumca 19(1971) 
473. Meliosma rigida Sieb. & Zurr. Abh. K. 
(Bayer.) Ak. Wiss. M. Ph. Kl. Munchcn 4, 2 (1845) 
153; Mio. Ann. Mus. Hot. Lugd.-Bat. 3 (1867) 93; 
Cat. Mus. Bot. I (1870) 23, incl. var. angusli/ulia 

MiQ., nomen; Maxim. Bot. Jahrb. 6 (1884) 60; 
Forbes&Hemsley, J. Linn.Soc. Bot. 23 (1886) 145, 
p.p., excl. M. pungens; Dunn, J. Linn. Soc. Bot. 38 
(1908) 358; Hayata, Ic. PI. Formos. 1 (1911) 161 
Dunn & Tutch. Kew Bull. Add. Ser. 10 (1912) 68 
Chun, Sunyatsenia 1 (1933) 180; Hand.-Mazz 
Beih. Bot. Centralbl. 52(1934) 166; Kaneh. Formos 
Trees ed. 2 (1936) 416, f. 372; Cufod. Oest. Bot. Z 
88 (1939) 267, incl. var. patens; Hara, Enum 
Sperm. Japon. 3 (1954) 121; Making, III. Fl. Jap 
(1954) 348, f. 1044; Walker, Imp. Trees Ryukyu Is 
(1954)200, f. 121; How, Acta Phytotax. Sin. 3 (1955) 
444; Gagnep. & Vidal, Fl. Camb. Laos & Vietnam 
1 (1960) 47, in obs.; Liu, III. Lign. PI. Taiwan 2 
(1962) 925, f. 762; Li, Woody Fl. Taiwan (1963) 503; 
Ohwi, Fl. Japan (1965) 613. - Quercus jama-buwa 
SiEB. in sched. ex Miq. Ann. Mus. Bot. Lugd.-Bat. 
3 (1867) 93, nom. inval. - Meliosma pungens auct. 
non (W. & A.) Walp.: Hook./. Fl. Brit. India 2 
(1876) 4, p.p., quoad pi. Japon. - Melio.sma patens 
Hkmsley ex Forbes &. Hemsley, J. Linn. Soc. Bot. 
23 (1886) 145. Meliosma harmandiana Pierre, 
II. For. Cochinch. 5 (1897) I. 360. - Meliosma 
gtomerulata Rehu. & Wils. in Sarg., PI. Wils. 2 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 10. Meliosma simplicifolia Walp. ssp. fruticosa (Blume) Beus. a. Fruiting twig, xO.5; b. detail of leaf 
undersurface, x2.5; c. endocarp, in different positions, x2.5 (a-c Kadim & Noor 395). 


Sabiaceae (van Beusekom & van de Water) 


(1914) 203. - Meliosma loheri Merr. Philip. J. Sc. 
10(1915) Bot. 38; Enum. Philip. Fl. PI. 2 (1923) 517. 
- Meliosma pannosa Hand.-Mazz. Anz. Ak. Wiss. 
Wien M.-N. Kl. 58 (1921) 179; How, Acta Phytotax. 
Sin. 3 (1955) 442; Gagnep. & ViDAt, Fl. Camb. Laos 
& Vietnam 1 (1960) 50, in obs., p.p. - Meliosma 
costata CuFOD. Oest. Bot. Z. 88 (1939) 266; How, 
Acta Phuotax. Sin. 3 (1955)444; Gagnep. & Vidai, 
Fl. Camb. Laos & Vietnam 1 (1960) 45; Vidax, Not. 
Syst. 16 (1960) 304. - Meliosma evrardii Gagnep. 
Not. Syst. 14(1952) 212, p.p. 

Leaves usually obovate-oblong to obovate-lanceo- 
late, sometimes oblong to lanceolate, 4-25(-32) by 
1.5-8(-ll) cm, base long-cuneate to acute, apex 
acute to cuspidate, without domatia; nerves 7-19 
pairs. Panicles lax to rather dense, 10-30 cm, 
branched up to the 2nd (3rd) order; axes more or less 
tomentose, sometimes woolly-pubescent, the pri- 
mary ones only exceptionally subtended by small 
leaves. Flowers more or less crowded, usually in 
dense glomerules, sessile; mature buds 1.7-2.2 mm 
diam. Sepals 5 (4). Inner petals 0.6-O.S mm, usually 
less than halfway bifid, lobes hardly or not 
divergent, rather narrow, fimbriate or ciliolate at the 
tips. Style about (1 .5-)2 times as long as ovary. En- 
docarps (sub)globose, not or not much oblique, 
(3.5-)4-5 mm diam., with fine reticulum; median 
keel blunt to rather sharp, hardly to distinctly promi- 
nent, often at one end running out into a minute ven- 
tral processus or tubercle; ventral pore not sunken, 
often somewhat spouted. 

Distr. Widely distributed in continental SE. 
Asia, incl. China, Laos, S. Vietnam (only at Hue), 
Taiwan (incl. Pescadores), Ryu Kyu Islands, Japan; 
in Malesia: Philippines (Luzon: Mountain Provi- 
nce). Fig. 9. 

E c o I . In evergreen broad-leaved or laurophyllous 
forests, on different soils; in dry as well as in wet 
places; altitude usually 100-1000 m, but in Luzon 
reported from 1200- 16(X) m. 

Field notes. Bark grey, smooth. Branches 
brown. Leaves lustrous green above, sometimes 
glaucous beneath. Fruit blue-purple to purplish 

Vern. Philippines: gahatan. If., Luzon; lasuit, 
Bondoc dial. 

Notes. Ssp. rigida is variable in quite some 
characters in its area outside Luzon, for instance in 
the degree of pubescence, leaf shape, and dentation. 
In continental SE. Asia the area of ssp. rigida bor- 
ders on or overlaps the areas of five or six other 
subspecies of M. simplici/olia, which substantially 
adds to the chance of confusing them, several 
specimens being hybrids, it is probable that these 
subspecies are ecologically isolated to a large extent 
and thus contact between them is prevented. 

Quercus gilva var. prucera Blumf:, Mus. Bot. 

Lugd.-Bat. 1 (1850) 306, was included in the synony- 
my of M. rigida by Hara, I.e., but I found it to 
belong to Quercus gilva Blume. 

c. ssp. fruticosa (Blume) Beus. Blumea 19 (1971) 
477, f. 28. - Meliosma fruticosa Blume, Rumphia 3 
(1849) 198; Walp. Ann. 2 (1852) 224; MiQ. Fl. Ind. 
Bat. 1,2 (1859) 614; Illustr. (1871) 73; K. & V. Bijdr. 
9 (1903) 133; Koord. Exk. Fl. Java 2 (1912) 545. - 
Meliosma elliptica Hook./. Fl. Brit. India 2 (1876) 5, 
p.p., excl. Sabia floribunda Miq.; King, J. As. Soc. 
Beng. 65, ii (1896) 456; Ridley, J. Str. Br. Roy. As. 
Soc. n. 33 (1900) 67; Fl. Mai. Pen. 1 (1922) 514; 
BuRK. & Hend. Card. Bull. S. S. 3 (1925) 364. - 
Meliosma lancifolia Hook./. Fl. Brit. India 2 (1876) 
5; King, J. As. Soc. Beng. 65, ii (1896) 456; Ridley, 
Fl. Mai. Pen. 1 (1922) 514. - Meliosma monophylla 
Merr. Philip. J. Sc. 4 (1909) Bot. 286; Enum. Philip. 
Fl. PI. 2 (1923) 517; Vidal, Not. Syst. 16 (1960) 306. 
- Fig. 8c, 10. 

Leaves usually oblong to lanceolate, 5-40(-45) by 
2-15 cm, base acute, apex acute to acuminate, 
densely pubescent on midrib and sometimes on 
nerves and lamina, beneath glabrous to tomentose, 
sometimes with domatia; nerves 7-25 pairs. Panicles 
usually lax, sometimes more dense, 10-50 cm, 
branched up to the 3rd (4th) order; eixes pubescent to 
short-tomentose, the lower primary ones subtended 
by small leaves or not. Flowers more or less crowded 
to solitary, (sub)sessile; mature buds 1.5-2 mm 
diam. Sepals 5 (4). Inner petals c. 0.7 mm, about 
halfway or somewhat less bifid; lobes divergent or 
not, usually rather narrow, more or less fimbriate, 
sometimes entire. Style about as long as ovary or 
shorter. Endocarps globose or ± ellipsoid, 
(4.5-)5.5-8 mm diam., with rather wide, coarse 
reticulum; median keel prominent, at one end often 
running out into a minute ventral processus; ventral 
pore not or not much sunken, not spouted. 

Distr. S. Peninsular Thailand (Sural) and 
Taiwan; in Malesia: common in the Malay Peninsula 
and Sumatra; W. Java, Lesser Sunda Islands (Sum- 
bawa, Flores), Borneo (Central Sarawak, Kinabalu, 
W. Kutai), and the Philippines (Luzon). Fig. 9. 

Ecol. Primary rain-forest, on various soil types, 
reported to occur on limestone, sand, volcanic loam, 
and andesite; altitude from sea-level up to 24(X) m. 

Field notes. Bark smooth, grey to brown, len- 
ticellatc, paperthin. Inner bark pale brown to dark 
brownrcd. Wood reddish to rcdbrown. Fruit yellow 
to pale red when ripening, dark red to brown when 

Vern. (all once noted). Malay Peninsula: bua 
palu, Sclangor, medang kerkulu, mengading, Malac- 
ca; Sumatra: lelagan, Gajo lang., Aceh, kaju djarap, 
k. gasir, k. si raga, Asahan, kaju ardong ardong, 
Toba, kabung kabung, Tapanuli, masadih pajo. 


Flora Malesiana 

[ser. I, vol. 10^ 

Simalur, kendung, Palembang, redjang, Djambi; 
Java: ki (iwu, Preanger; Flores: kaju sar, Philip- 
pines: malaligas. Tag. 

d. spp. pungens (Wall, ex W. & A.) Beus. Blumea 
19 (1971) 466. - Millingtonia pungens Wall, ex 
W. & A. Edinb. New Phil. J. 15 (1833) 178; Prod. 1 
(1834) 115; Wight, Ic. 3 (1845) t. 964/3. - Meliosma 
pungens (Wall, ex W. & A.) Walp. Rep. 1 (1842) 
423; Ann. 1 (1848) 135; Thw. Enum. PI. Zeyl. (1858) 
59; Bedd. F1. Sylv. 3 (1871) 77; ibid. t. 160; Merr. 
Contr. Am. Arb. 8 (1934) 94; Vidal, Not. Syst. 16 
( 1 960) 306. - Meliosma wightii Planch, ex Brandis, 
For. Fl. (1874) 116;HooK./. Fl. Brit. India 2 (1876) 
4. - Fig. 8a. 

Leaves elliptic to oblong, sometimes lanceolate, 
5-20(-30) by 2-8(-10) cm, without or with some 
distant teeth, acute to rounded at the base, acute to 
acuminate at the apex, usually distinctly pubescent 
on midrib and sometimes on nerves above, sparsely 
to moderately pubescent beneath especially on mid- 
rib and nerves, usually with domatia; nerves 7-18 
pairs. Panicles lax to dense, (5-) 10-55 cm, branched 
up to the 2nd (3rd) order; axes rather coarse, densely 
short-tomentose, the lower primary ones almost 
always subtended by small leaves. Flowers crowded 
in dense glomerules, sessile; mature buds 2-2.5 mm 
diam. Sepals {8-)9-ll(-\i). Inner petals c. 1 mm, 
slightly bifid; lobes divergent, wide, glabrous. Style 
about as long as ovary. Endocarps (sub)globose, 
often rather irregular, 3.5-5.5 mm diam., with 
usually lax reticulum; median keel distinct but not 
very prominent, not running out into a ventral pro- 
cessus; ventral pore hardly or not sunken, not spout- 

Distr. Sri Lanka and Deccan Peninsula; in 
Malesia: N. Sumatra (Gajo Lands, Takengon), one 
collection. Fig. 9. 

Ecol. Mountain forest, 1500-2000 m altitude. 

3. Meliosma sumatrana (Jack) Walp. Ann. 1 (1848) 
135; MiQ. Fl. Ind. Bat. 1, 2 (1859) 617; Sum. (1861) 
203; lUustr. (1871) 75; Hook./. Fl. Brit. India 2 
(1876) 6; KooRD. Minah. (1898) 408; Suppl. Cel. 2 
(1922) 7, t. 56; ibid. 2 (1922) 28; Merr. Enum. Born. 
(1921) 363; Enum. Philip. Fl. PI. 2(1923) 518; Contr. 
Arn. Arb. 8 (1934) 95; Merr. & Perry, J. Am. Arb. 
20(1939) 357; van Beusekom, Blumea 19 (1971) 485. 
- Millingtonia sumatrana Jack, Mai. Misc. 2 (7) 

(1822) 30; Hook. J. Bot. 1 (1834) 378; Merr. J. Arn. 
Arb. 33 (1952) 236. - Meliosma nitida Blume, Cat. 

(1823) 32; Nees, Flora 8 (1825) 106; Hassk. Tijd. 
Nat. Gesch. Phys. 10 (1843) 139; Cat. Hort. Bog. 
(1844) 226; Blume, Rumphia 3 (1849) 202, t. 169, 
incl. var. tridenta Blume, var. cerasiformis Blume et 
var. splendens Blume; Walp. Ann. 2 (1852) 225; 
Miq. Fl. Ind. Bat. 1, 2 (1859) 617; Sum. (1861) 203, 

520; Illustr. (1871) 74; King, J. As. Soc. Beng. 65, ii 
(1896) 457; K. & V. Bijdr. 9 (1903) 117; Koord. Exk. 
Fl. Java 2 (1912) 546, f. 81; Atlas 2 (1914) 377; 
Ridley, Fl. Mai. Pen. 1 (1922) 515; Baker/, in Ren- 
dle, J. Bot. 62 (1924) Suppl. 30; Burk. & Hend. 
Card. Bull. Str. Settl. 3 (1925) 364; Heyne, Nutt. PI. 
(1927) 1002; Merr. & Perry, J. Arn. Arb. 20 (1939) 
357; Backer & Bakh./. Fl. Java 2 (1965) 145. - Irina 
integerrima Blume, Bijdr. (1825) 231, non Hassk. 
PI. Jav. Rar. (1848) 284 ('Iriney, Walp. Rep. 1 
(1849) 416; Blume, Rumphia 3 (1849) 202, in syn. 
sub M. nitida. - Millingtonia nitida Schult. & 
ScHULT. Syst. Veg. Mant. 3, add. 2 (1827) 250; 
DiETR. Syn. PI. 1 (1839) 103. - Meliosma confusa 
Blume, Rumphia 3 (1849) 200; Walp. Ann. 2 (1852) 
225; Miq. Fl. Ind. Bat. 1, 2 (1859) 616; Sum. (1861) 
203, 520; Illustr. (1871) 74. - Meliosma cuspidata 
Blume, Rumphia 3 (1849) 202; Miq. FI. Ind. Bat. 1, 
2 (1859) 617; Illustr. (1871) 74; Hall./ Meded. 
Rijksherb. 1 (1910) 2; Merr. Enum. Born. (1921) 
362. - Meliosma pinnata (non Walp.) Koord. 
Minah. (1898)408. - Meliosma diepenhorstii Valet. 
Ic. Bog. 2 (1904) 195, t. 150. - Meliosma elmeri 
Merr. PI. Elm. Born. (1929) 177. - Meliosma 
philippinensis Merr. & Perry, J. Arn. Arb. 20 
(1939) 357. 

Evergreen tree, up to 15-20(-25) m. Leaves 
2-5(-6)-jugate; rachis terete, 6-50 cm, including 
the up to c. 25(-30) cm long petiole, up to c. 10(- 15) 
mm across, rarely slightly pubescent, usually with 
distinctly swollen base; leaflets usually elliptic to 
lanceolate, (3-)5-35(-50) by (1.5-)2.5-I5(-20) 
cm, base cuneate to rounded, shortly narrowed into 
the petiole, apex acuminate to caudate, usually en- 
tire, beneath rarely more or less pubescent, without 
domatia; midrib slightly prominent above; nerves 
(5-)7-13(-19) pairs, ascending, nearly always 
looped and joined; petiolules very short or up to c. 
6 cm, usually distinctly swollen at the base especially 
in older leaves. Panicles usually terminal, usually 
narrowly, sometimes widely pyramidal, 7-50(-75) 
cm, usually profusely branched up to the 4th order, 
rather stiff and coarse, puberulous, bearing numer- 
ous crowded flowers; primary side-axes usually 
rather short, up to c. 30 cm, the lower ones excep- 
tionally subtended by small to reduced leaves; bracts 
ovate to narrowly triangular, up to c. 6 mm, ± 
puberulous. Pedicels absent or short, up to c. 2 mm. 
Mature buds (1 .5-)2-3(-3.5) mm diam. Sepals 5 or 
4, ovate, unequal, the inner 3 or 4 c. 1-2 mm, the 
outer 1 or 2 usually smaller, often minute, sometimes 
lowered on the pedicel, sometimes puberulous out- 
side, especially the outer ones, with entire or 2- or 3- 
lobed, often ciliolate margin. Outer petals glabrous. 
Inner petals elliptic to lanceolate or strap-shaped 
with wide-truncate tip, (1.2-)1.5-2(-3) mm, acute 
to slightly bifid or retuse and frayed at the tip. Ovary 


Sabiaceae (van Beusekom & van de Water) 


Fig. 11. Generalized areas of Meliosma sumatrana (Jack) Walp. (thick line) and M. lanceolata Blume (thin 

line); the small ova! areas indicate the localities of M. lanceolata var. polyptera (Miq.) Beus. The distribution 

of M. hirsuta Blume is indicated by a dot, that of M. rufo-pilosa Hend. by squares. 

0.5-1 mm, glabrous. Fruit globose to short-ellip- 
soid, when ripe 1-3 cm diam., with rather thick 
spongy to pulpy mesocarp; endocarp ellipsoid, some- 
times nearly globose, 0.7-2 cm diam., with almost 
smooth to somewhat lumpy surface, often with a few 
faint to sharply prominent ribs; median keel distinct, 
slightly elevated to sharply prominent, at one end 
often running out into a more or less prominent curv- 
ing, at the other end sometimes into a minute tuber- 
cle; ventral pore mostly rather wide, usually some- 
what sunken. 

Distr. Malesia: Sumatra (incl. Nias, Batu & 
Sipora Is., Banka), Malay Peninsula (incl. Penang 
I.), Anambas Is., W. half of Java, throughout Bor- 
neo, Sulawesi, and the Philippines (Mindanao, 
Palawan). Common. Fig. II. 

Lcol . Primary and secondary lowland and mon- 
tane rain-forest, up to c. 2200 m altitude. Found on 
various soils, fertile as well as infertile, in dry to wet 
localities, in dense to open forests, by streams as well 
as on hilltops and ridges. 

Field notes. Often a crooked tree, irregularly 
branched. Trunk sometimes with small buttresses. 
Bark surface grey to brown, smooth, with lenticels. 

often with shallow fissures, sometimes said to be 
dimpled, patchy or scaly. Inner bark 0.5- 1 cm thick, 
soft, fibrous, light yellow or dirty white, soon turn- 
ing pink, brownish, reddish, or rusty after exposure. 
Sapwood said to be whitish, yellowish, creamy 
orange, or brownish. Sap without special smell or 
taste. Leaves bright green on both sides. Flower col- 
our varying from white, cream, or greenish, to partly 
or entirely pinkish to red. Fruit first yellow, then 
yellow with red to red when ripe; pulp white, turning 
quickly blood-red on exposure, finally becoming 
black, sweetish to tasteless. 

Uses. The species was proposed by Koorders & 
Valeton, I.e., for reforestation purposes. In Min- 
danao the triturated bark and leaves are several times 
reported to be in use as a medicine applied for 
wounds, to soothe itchy skin or - charred and put in 
water - against tympanites. It was also .said to be 
used in agricultural rituals. The fruits are many times 
reported to be edible. 

Vern. Malay Peninsula: pa-on^, Saki name, and 
mennadinfi hesar, both from Pahang, huah mata 
ikan, Icmuan, Sclangor, pokoh haran, Negri Scm- 
bilan, kaju kahwa kanlu, membuloh, pokoh gra6 


Flora Malesiana 

[ser. I, vol. 10^ 

jantan, p. mala gajah, p. paigigi, p. pinang plandog, 
p. ravoa antoo, pudding utan, Malacca, pelanlu; 
Sumatra: laon, si paturut, sringkut, Karo country; 
kaju durung durung, k. ining ining, Tapianuli, tarn- 
pa bussie, Priaman, marazat, Mt Kerinci; Java: ki 
liwu (landuk) (bodas), S, ki huut, Udjong Kulon; 
Borneo, Sarawak: buliliap, Kenyah dial., maiak, 
Kayan dial., buiu manuk, Iban, bilonok, Dyak; 
Sabah: bung lai, Sungai, gapas gapas, kapas kapas, 
keriyan, Dusun, illulal, limpangol, tunjang, Murut; 
SE. Borneo: tambalilin, tandao, Dyak, Tidung dial.; 
djangkanggunung, Bandjar lang., Riom dial.; 
Sulawesi: see Koord. Minah. (1898) 408; eng- 
golokia, W. Toradja dial., pulu putu, situi, Tobela 
lang., Malili, pobumengo, Gorontalo; dama, Torai 
dial., Menado; Philippines: carabo-rabo, daborabo, 
kadabudabo, karabu-rabu, magobaylung, mahag- 
kol, yagabogan, Mbo, Buk., bentinguasay, gepulu, 
Zamboanga, waat, Cebuano, Mt Apo, salalab, 
Moro dial., garong, gimbingimbing. Sub., sumaga- 
sa, Bag. 

Notes. Meliosma sumatrana is very constant in 
its discriminative characters (especially the promi- 
nent midrib and entire inner petals), but there is 
nevertheless some geographical variation, especially 
in the northern part of Borneo (Sarawak, Sabah). As 
general tendencies may be noticed that towards the 
centre of the area leaflets and fruits increase in size 
and dentate leaflets become more common. More- 
over, the number of leaflet pairs decreases when the 
leaflets are larger. 

Sterile hybrids between M. sumatrana and M. pin- 
nata ssp. ridieyi are rarely found (Sabah). 

DocTERS VAN Leeuwen (Zoocecidia Neth. East 
Indies, 1926, 339, f. 612) described a leaf-gall on a 
specimen from Sulawesi. This type of galls (usually 
ball-shaped, c. 4 mm, ending in a short mucro, and 
surrounded by a calyx-like circumvallation) is rather 
commonly met with in this species, not only in 
specimens from Sulawesi, but also from Borneo, 
Sumatra, and the Malay Peninsula. The galls do not 
only occur on the lower surface of the leaflets, but 
occur also on the upper surface, and on rachis and 
petiolules, often very many crowded together. 

4. Meliosma lanceolata Blume, Cat. (1823) 32; Nees, 
Flora 8 (1825) 106; Hassk. Cat. Hort. Bog. (1844) 
226; Blume, Rumphia 3 (1849) 200, t. \6i,p.p., incl. 
var. pendula Blume, var. membranacea Blume, var. 
chart acea Blume et var. obliqua Blume; Walp. Ann. 
2 (1852) 224; Miq. F1. Ind. Bat. 1 , 2 (1859) 614; Sum. 
(1861) 203, 520; lllustr. (1871) 74, p.p.; Hook./. F1. 
Brit. India 2 (1876) 7; King, J. As. Soc. Beng. 65, ii 
(1896) 458; Ridley, J. Str. Br. Roy. As. Soc. n. 33 
(1900) 67; K. & V. Bijdr. 9 (1903) 125; Hall./. Med. 
Rijksherb. 1 (1910) 2, in obs.; Koord. Exk. Fl. Java 

2 (1912) 546; Merr. Enum. Born. (1921) 363; 
Ridley, Fl. Mai. Pen. 1 (1922) 516, f. 51; Baker/. 
in Rendle, J. Bot. 62 (1924) Suppl. 30; Craib, Fl. 
Siam. Enum. 1 (1926) 340; Ridley, Kew Bull. (1926) 
63; Merr. PI. Elm. Born. (1929) 176; Hochr. Can- 
dollea 6 (1936) 467, incl. var. genuina Hochr.; 
Backer & Bakh./ Fl. Java 2 (1965) 145; van 
Beusekom, Blumea 19 (1971) 489. - Millingtonia 
lanceolata Schult. & Schult. Syst. Veg. Mant. 3, 
add. 2 (1827) 250; Dietr. Syn. PI. 1 (1839) 103. - 
Meliosma polyptera Miq. Sum. (1861) 203, 520; 
lllustr. (1871)73. - Meliosma levis KwG, J. As. Soc. 
Beng. 65, ii (1896) 457; Ridley, Fl. Mai. Pen. 1 
(1922) 515. - Meliosma nervosa K. & V. Bijdr. 9 
(1903) 129; Koord. Exk. Fl. Java 2 (1912) 546; Atlas 
2 (1914) t. 376; Fl. Tjibodas 2 (1923) 158; Merr. & 
Perry, J. Arn. Arb. 20 (1939) 359, in obs.; Backer 
& Bakh./ Fl. Java 2 (1965) 145. 

Evergreen tree, up to c. 25(-30) m. Twigs often 
with conspicuous leaf-scars. Leaves (3-)7-18(-25)- 
jugate; rachis terete, (10-)30-100 cm, including the 
5-30 cm long petiole, up to c. 8 mm diam., usually 
with distinctly swollen base, usually ± lenticellate; 
leaflets usually oblong to lanceolate, hardly or not 
asymmetrical, 5-20 by 2-7 cm, not or only slightly 
increasing in size towards the top of the leaf, often 
the lowermost pairs much smaller, base usually acute 
to rounded, apex acuminate to cuspidate, glabrous 
to moderately pubescent, always without domatia; 
midrib usually deeply impressed above; nerves 5-16 
pairs, ascending, looped. Panicles terminal, nearly 
always pendulous and lax, rarely erect (then also 
small), pyramidal, usually large, (15-)50-150 cm 
and profusely branched up to the 3rd order, ± pu- 
bescent, bearing numerous glomerulate or crowded 
flowers which are usually spicately arranged, the 
glomerules often with regular space; main axis terete, 
often bent down abruptly at the base; primary side- 
axes many, usually long, up to c. 90 cm, never 
subtended by leaves; bracts ovate to narrowly 
triangular, up to c. 5 mm, ± pubescent. Pedicels ab- 
sent, up to c. 1 mm. Mature buds 1.5-2 mm diam. 
Sepals 5 (4), ovate, more or less unequal, the inner 3 
or 4 c. 1 mm, the outer 2 or 1 usually much smaller, 
often minute and sometimes slightly keeled, 
sometimes somewhat lowered on the pedicel, all 
glabrous, and with an entire margin. Outer petals 
1.5-2 mm. Inner petals about halfway bifid, c. 0.6 
mm, with ciliolate, rarely glabrous lobes, usually 
with a minute central lobule. Filaments c. 1 mm. 
Ovary (0.5-)0.7(-l) mm, usually densely, some- 
times sparsely pubescent, rarely glabrous. Fruit 
(sub)globose, when ripe 7-10 mm diam.; endocarp 
subglobose, often somewhat depressed to applanate 
at the ventral side, usually strongly obUque, (5-)6-9 
mm diam., with usually distinct, rather coarse, most- 
ly sharply prominent reticulum; median keel sharp 


Sabiaceae (van Beusekom & van de Water) 


and prominent, at one end often running out into a 
small to minute ventral processus or tubercle; ventral 
pore not or not much sunken. 

D i s t r . Nicobar Is. , extreme South of Peninsular 
Thailand; in Sfalesia: Sumatra (incl. Simalur, Batu, 
and Banka Is.), W. Java, Borneo (northern halO- 
Not uncommon, scarce in Borneo. Fig. 11. 

Ecol. Primary and often secondary forests, at 
low and medium altitudes, occasionally ascending to 
1500 m,/. nervosa to 2900 m, on various soil types. 

Field notes. Outer bark grey to brown, rather 
smooth, later with longitudinal cracks, thin, often 
lenticellate. Inner bark 0.5-1 cm, several times said 
to be (light) red, orange brown, or redbrown, also 
dirty white and then turning rusty after e.xposure. 
Wood soft, white or pale yellow to light yellow 
brown. Crown low, irregular and lax, with few usual- 
ly crooked branches. The conspicuous large leaves 
are rather crowded at the end of the twigs. Leaflets 
when young red-brownish. Flowers white or yellow- 
ish to pink or red (sometimes different colours in the 
same panicle). Fruits, first dirty red, then bluish black 
when ripe. 

Vern. Malay Peninsula: medang siri, Malacca; 
Sumatra: kabung kabung (blumut), Batak lang., 
Simelungun dial., bulung manuk, Batak lang., Karo 
dial., sondang, sonlang, Timor on N. Sumatra, kaju 
buluk hudjan, Lampong, angke foluh pajo, silaora, 
sarin silo bulung, tulun surin or seulang (pajo), t. 
lungke ali, Simalur I.; W. Java: ki tiwu, S, often 
used as well for M. pinnata and M. sumatrana (also 
with the addition lalaki, mindi bodas or persawon), 
suren ieuweung, S. See also under var. lanceolata f. 
nervosa and var. polyptera. 

Notes. Meliosma lanceolata is generally very 
well characterized by its large pendulous panicles and 
its long leaves with many usually lanceolate leaflets. 
Nevertheless it shows a wide variation especially in 
number but also in shape and size of the leaflets and 
the panicles. On the islands west of Sumatra 
(Simalur, Nias, Batu) specimens are found with nor- 
mal inflorescences but only 3-5-pinnate leaves, and 
elliptic, sometimes subrotund, large leaflets. Transi- 
tions to this extreme are common. There is another 
deviating form, however, which takes a separate 
position. It has many small, mostly lanceolate leaf- 
lets which otherwise do not differ from those of M. 
lanceolata. Also the panicles agree with that species. 
In view of the wide variability in the leaves of M. 
lanceolata, I prefer to include it here and I have 
reduced it to a variety. The varieties and forms can 
be distmguished as follows: 

a. var. lanceolata. 

f.cavcs(3 )6 18 jugate, with up IOC. lOOcm long 
rachis (including the petiole); leaflets elliptic to 
lanceolate, mcdium-si/cd to large, 5 20( 25) by 

(2-)2.5-7(-10) cm, index (l.5-)2-5(-6), without 
or with teeth, glabrous or pubescent. 

Notes. In the lowland parts of its area var. 
lanceolata is nearly always very constant in the main 
characters. Mainly at higher elevations, however, 
forms occur which deviate considerably, often to 
such an extent that it is very difficult to separate them 
from less typical forms of the otherwise well distinct 
M. pinnata ssp. ferruginea and ssp. ridleyi; in a few 
cases, especially when the material is incomplete, this 
can only be done by a specialist who is thoroughly ac- 
quainted with habitus and variability of both species. 

For instance, a form with erect, unusually short 
panicles (sometimes only 15 cm long) and other de- 
viating characters may be met with. It occurs mainly 
in the montane zone; transitional forms are found 
lower, and these show a more or less gradual fading 
of typical lanceolata characters. Specimens of this 
mountain form have been described from Java by 
KooRDERS & Valeton, I.c, as M. nervosa. In my 
opinion this species should be reduced to the rank of 
a form only; see below. 

forma lanceolata. 

Leaves (3-)6-18-jugate, with elliptic to lanceo- 
late, glabrous to pubescent leaflets. Panicles pen- 
dulous, usually much longer than 50 cm. Inner petals 
ciliolate. Ovary pubescent. 

/ormo nervosa (K.&V.)Beus. Blumea 19(1971)493. 

- M. nervosa K. & V., vide supra. 

Leaves not more than 8(-10)-jugate, with usually 
elliptic glabrous leaflets. Panicles erect, shorter than 
c. 50cm, minimum length c. 15 cm. Inner petal most- 
ly glabrous. Ovary pubescent to glabrous. 

Distr. Malesia: Sumatra (G. Leuser, G. Talak- 
mau), W. Java. 

Ecol. Mountain forest, 1300-2900 m altitude. 
The tree can reach a height of 30 m by I m diam. 

Vern. Java: ki Ijermeh badak, ki tjermeh beu- 
reum, S. 

b. var. polyptera (Miq.) Beus. Blumea 19 (1971) 492. 

- M. polyptera Miq., vide supra. 

Leaves 12-25-jugate, with at most 50 cm long 
rachis (including the petiole); leaflets oblong to 
linear-lanceolate, small, 4-11 by 1-2 cm, entire, 

Distr. Malesia: Sumatra (Asahan, W. Coast). 
Fig. II. 

Ecol. At low altitudes. 

Vern. Sumatra: badar badar, Lubuk Alung, tan- 
dikat batu, Priaman, simarpapahu, Mula Padang. 

5. Meliosma hirsuta Blumh, Rumphia 3 (1849) 200; 
Waip. Ann. 2 (1852) 225; Mio. Fl. Ind. Bat. I, 2 
(1859) 616; Sum. (1961) 203; lllustr. (1871) 74; 


Flora Malesiana 

[ser. I, vol. 10^ 


Sabiaceae (van Beusekom & van de Water) 


Merr. Enum. Born. (1921) 363; van Beusekom, 
Blumea 19 (1971) 493. 

Evergreen small tree, c. 5 m. Leaves 15-20-(or 
probably more-)jugate; rachis 50-100 cm including 
the 10-20 cm long petiole, up to c. 6 mm across, 
more or less hirsute, usually with distinctly swollen 
base, sometimes sparsely lenticellate; leaflets (sub) 
sessile, those in medium and upper part of the leaf 
linear-lanceolate, 10-20(-25) by (1.5-)2-3 cm, in- 
dex 5-10, the lower ones (ovate-)lanceolate to ovate, 
gradually decreasing in length towards the base of 
the leaf, up to only c. 3 cm, base rounded to acute, 
sometimes slightly oblique, apex acuminate to 
caudate, with entire to remotely spinously dentate 
margin, thin-chartaceous, above glabrous except for 
some pubescence on the midrib, beneath moderately 
to sparsely hirsute especially on midrib and nerves, 
without domatia; midrib above flat to slightly im- 
pressed; nerves widely apart, (5-)8-12 pairs, as- 
cendmg, looped and joined into a distinct marginal 
nerve situated at 2-4 mm from the margin; venation 
distinct, wide, reticulate; petioles absent or up to c. 
1 mm, terminal one often longer, up to c. 8 mm, 
densely hirsute, not swollen at the base. Panicles and 
flowers as in typical M. lanceolala, but sepals up to 
c. 1.5 mm. Fruit as in M. lanceolala. 

Distr. Malesia: Sumatra (West Coast: G. Malin- 
tang), only one collection. Fig. 11. 

Notes. This species was by Blume erroneously 
recorded to occur in S. Borneo. 

Meliosma hirsuta is doubtless very closely related 
to M. lanceolala, but very well distinct by its leaf 

See also Pimela angustifolia under the dubious 

6. Meliosma pinnata (Roxb.) Maxim. Bull. Ac. Imp. 
Sc. St. Petersb. 12 (1867) 64; Melanges 6: 263. - 
Millingtonia pinnata Roxb. F1. Ind. 1 (1820) 103. - 
Fig. 12. 

For further synonyms, see under the subspecies; 
for a complete synonymy, see van Beusekom (1 97 1 : 

Evergreen, sometimes deciduous tree, small to up 
to c. 42 m. Twigs often with conspicuous leaf-scars. 
Leaves 2- II -jugate; rachis terete, (2-)5-40(-60) 
cm, including the up to c. 15(-25) cm long petiole; 
leaflets usually ovate, elliptic, or obovate to ovate- 
oblong, sometimes lanceolate, often asymmetric. 

1.5-25 by l-IO cm, usually increasing in size to- 
wards the top of the leaf, base usually acute to 
rounded, rarely slightly emarginate, apex acuminate 
to cuspidate, entire or dentate, usually slightly to 
densely pubescent, often with domatia; midrib flat to 
impressed above; nerves 3-15 pairs, ascending, 
looped; petiolules up to 5 cm, terminal one usually 
longest, not or not much swollen at the base. Panicles 
terminal, erect, sometimes somewhat pendulous, 
dense to lax, widely to narrowly pyramidal, 
10-55(-70) cm, usually profusely branched up to 
the 4th order, bearing numerous solitary to usually 
crowded flowers; primary side-axes usually many, up 
to 35(-60) cm, lower ones sometimes subtended by 
small to reduced leaves; bracts ovate to narrowly 
triangular, up to c. 5(-10) mm, more or less pubes- 
cent. Pedicels absent or up to 3(-4) mm. Mature 
buds (1.5-)2(-3) mm diam. Sepals 5 or 4, ovate, 
unequal, the 3 or 4 inner ones 1-1.5 mm, the outer 
1 or 2 usually smaller, often minute, sometimes 
lowered on the pedicel, sometimes slightly keeled, 
glabrous or pubescent outside, all entire, usually 
ciliolate. Outer petals usually glabrous. Inner petals 
more or less deeply bifid, (0.3-)0.6(-l) mm, gla- 
brous, ciliolate or fimbriate at the tips, often with a 
minute central lobule, often frayed at the tips. 
Filaments c. 1 mm. Fruit (sub)globose to obovoid, 
when ripe (3-)4-10(-ll) mm diam., with thin 
mesocarp; endocarp (sub)globose, oblique or not, 
(2.5-)3.5-9(-10) mm diam., with more or less pro- 
minently reticulate surface; median keel usually 
distinct and more or less prominent, at one end 
sometimes running out into a small to minute pro- 
cessus or tubercle, and sometimes curving outwards 
at the other end; ventral pore usually rather narrow, 
whether or not sunken. 

Distr. Throughout SE. Asia, from Sri Lanka and 
China to Japan; throughout Malesia as far as New 
Guinea (incl. New Britain). Fig. 13. 

E c o 1 . Forests under moist tropical to subtropical, 
sometimes warm-temperate conditions, on various 
soils, from sea-level up to c. 3000 m altitude. 

Notes. Meliosma pinnata covers a very large area 
in which it has developed a complex and wide varia- 
tion pattern. It can be divided up into nine well- 
marked subspecies. Four of these are widely distrib- 
uted, whereas five have a limited distribution. The 
first group, the subspecies arnottiana, ridleyi, 
macrophylla aind ferruginea , are considered primary 

Fig. 12. Meliosma pinnata (Roxb.) Walp. s.sp. macrophylla (Murr.) Beus. a. Flowering twig, xO.33; b. half- 
opened flower, y 5; c. outer petal with adhering staminodc; d. flower with outer petals removed and stamens 
snapped backward; e g. stamen with adhering inner petal, in different positions; h. pistil with surrounding 
disk; (. ovary, length section, all x IO;y. ripe fruit, x 3; *-/. endocarp in different positions, x 3 (a / Sulit 

F'NH 32941 . j I Kostermans 691 1 ). 


Flora Malesiana 

[ser. I, vol. 10^ 

Fig. 13. Generalized areas of the subspecies of Meliosma pinnata (Roxb.) Walp., and distribution of M. 

sarawakensis Ridley. 

subspecies; they centre in W. Malesia. The sub- 
species of the second group occur scattered at the 
periphery of the area of M. pinnata; I consider them 
secondary offsplits from the primary subspecies, viz. 
ssp. pinnata and ssp. angustifolia (Merr.) Reus. 
from ssp. arnottiana, and ssp. pendula, ssp. sylva- 
tica, and ssp. humilis from ssp. macrophylla. 

The areas of the secondary subspecies fall partly or 
entirely within the area of the primary subspecies 
from which they are derived, but they are ecological- 
ly isolated from these, usually by preference for dif- 
ferent altitudinal zones; transitional or hybrid forms 
are sometimes found. The areas of the four primary 
subspecies, on the other hand, all touch or only 
slightly overlap mutually, but generally they are per- 
fectly replacing, and usually there is also different 
ecological preference. Due to the scarcity of collec- 
tions from critical regions, especially Sumatra, 
Borneo and Sulawesi, it is mostly not clear how the 
relation is in contact zones. There is some evidence 
that one or two mutually may behave as good 
species, where one or two others may be connected 
by transitional forms, but in general the evidence re- 

quired is still wanting. In this respect the picture is 
not so complete as it is in M. simplicifolia. 
The type subspecies does not occur in Malesia. 


1. Ovary glabrous or only with a few hairs. Sepals 

and petals always glabrous. 
2. Leaves 3-5-jugate; leaflets dentate (sometimes 
only a few teeth), with domatia in the axils of the 
nerves beneath which are sometimes obscured by 
very dense tomentum of the leaf-blade 

g. ssp. humilis 
2. Leaves (3-)4-6(-7)-jugate; leaflets dentate or 
not, without domatia, never with very dense 
3. Leaflets entire, index (l-)1.5-3, mostly round- 
ed or obtuse to truncate or emarginate at the 
base. Medium-sized to large trees 

d. ssp. macrophylla 
3. Leaflets dentate (sometimes very sparsely), in- 
dex (l-)1.5-4(-5), acute or rounded, obtuse, 
truncate or emarginate at the base. Small to 
medium-sized trees, rarely shrubs. 


Sabiaceae (van Beusekom & van de Water) 


4. Leaflets moderately to rather densely villous- 
pubescent (often more or less glabrescent when 
older), mostly (especially lower ones) rounded 
to truncate at the base, index 1.5-3(-4). En- 
docarps 6-7 mm diam., without ventral pro- 
cessus. Above c. 1800 m alt. e. ssp. pendula 
4. Leaflets sparsely to densely short-pubescent, 
rarely subglabrous, mostly with acute base, in- 
dex I.5-4(-5). Endocarps 5-7.5 mm diam., 
mostly with a small but distinct ventral pro- 
cessus. Below c. 1000 m altitude. 
5. Leaves (3-)4-6(-7)-jugate; leaflets acute at 

the base f . ssp. sylvatica 

5. Leaves 3-5(-6)-jugate; lateral leaflets 
rounded to truncate at the base 

d. ssp. macrophylla (Celebes form) 
1. Ovary entirely, rarely partly, but always densely 
pubescent. Sepals and petals glabrous or pubes- 
6. Sepals and usually also outer petals moderately 
to densely pubescent on the outside. Leaflets en- 
tire, index (l-)1.5-3 c. ssp. ferruginea 

6. Sepals and petals glabrous or rarely a few hairs 
on the outer sepals only. Leaflets entire or den- 
tate, index (1-)1 .5-4(-5). 
7. Endocarps 4.5-9(-10) mm diam., usually with 
more or less sunken ventral pore. Inner petals 
with fimbriate or ciliolate, rarely glabrous 
lobes. Leaflets never with domatia 

b. ssp. ridleyi 

7. Endocarps (2.5-)3-4.5 mm diam., not with 

sunken ventral pore. Inner petals with usually 

glabrous, sometimes at the tips ciliolate or 

frayed lobes. Leaflets with or without domatia 

a. ssp. arnottiana 

a. ssp. arnottiana (Wight) Beus. Blumea 19 (1971) 
499. - Sapindus ? microcarpus W. & A. Prod. 1 
(1834) 112, nom. illeg., non R. & P. (1804); Wight, 
III. Ind. Bot. 1 (1840) 142; Walp. Rep. 1 (1842)416, 
423. - Millingtonia arnottiana Wight, III. Ind. Bot. 
1 (1840) 144, t. 53. - iVellinglonia arnottiana Meiss . 
PI. Vase. Gen. (Comm.) 2 (1840) 207, in nolo. - 
Millingtonia samhucina Jungh. Tijd. Nat. Gesch 
Phys. 8 (1841) 365. - Meliosma arnottiana Walp 
Rep. I (1842) 423; Thw. Enum. PI. Zcyl. (1858) 59 
Bedd. FI. Sylv. 3 (1871)77; ibid. t. 160; Hook./. FI 
Brit. India 2 (1876) 6; Trim. FI. Ceyl. 1 (1893) 315 
Bra.sdis, Indian Trees (1906) 195; Gamble, FI. Pres 
.Madras I (1918) 256. - Meliosma glauca Blume 
Rumphia 3 (1849) 200. I. I68B, nom. illeg. \ Walp 
Ann. 2 (1852) 225; Hassk. Hon. Bog. I (1858) 140 
MiO. M. Ind. Bat. I, 2 (1859) 615; K. & V. Bijdr. 9 
(1903) 135. ind. var. floribunda (Biijmi.) K. & V. 
•Hall./. Mcdcd. Rijkshcrb. I (1910) 2; KrK)Rn. Fxk 
n. Java 2 (1912) 546; M. Tjibodas 2 (1923) 157 
Baker/ in Rcndlc. J. Bot. 62 (1924) Suppl. 30. 

Meliosma floribunda Blume, Rumphia 3 (1849) 200; 
Walp. Ann. 2 (1852) 225; MiQ. FI. Ind. Bat. 1, 2 
(1859) 615; lllustr. (1871) 74; K. & V. Bijdr. 9 (1903) 
137; Hall./ Meded. Rijksherb. 1 (1910) 2; Koord. 
Exk. FI. Java 2 (1912) 546. - Meliosma sambucina 
MiQ. lllustr. (1871) 74; K. & V. Bijdr. 9 (1903) 137, 
in obs. - Meliosma luzonensis Merr. Publ. Govt. 
Lab. Philip. 29 (1905) 24; Elmer, Leafl. Philip. Bot. 
2 (1908) 492, in obs. ('luzonica'); Merr. Enum. 
Philip. FI. PI. 2 (1923) 517. - Meliosma multiflora 
Merr. Publ. Govt. Lab. Philip. 29 (1905) 25; Enum. 
Philip. FI. PI. 2 (1923) 517. - Melliosma ferruginea 
(non Blume) Koord. Gedenkb. Jungh. (1910) 177. - 
Meliosma apoensis Elmer, Leafl. Philip. Bot. 10 
(1939) 3784, descr. angl. - Meliosma cannarioides 
Elmer, Leafl. Philip. Bot. 10 (1939) 3785, descr. 
angl. - Meliosma ferruginea (non Blume) Backer & 
Bakh./ FI. Java 2 (1965) \45, p.p., quoad M. glauca 
et floribunda. 

Small to medium-sized, rarely big tree, up to c. 
20(-30) m. Leaves (2-)3-7(-8)-jugate; leaflets 
ovate to ovate-oblong, elliptic, or lanceolate, small 
to up to c. 25 by 10 cm, index (l-)1.5-4(-5), acute 
to truncate at base, entire or dentate, chartaceous to 
coriaceous, often with domatia. Panicles erect, 
spreading, lax to dense, lower primary side-axes 
usually subtended by small or reduced leaves. Sepals 
glabrous or the outer ones rarely with a few hairs. 
Petals glabrous, inner ones sometimes a bit ciliolate 
or frayed at the tips of the lobes. Ovary densely 
pubescent, very rarely subglabrous. Endocarps (sub) 
globose, not or not much depressed, hardly or not 
oblique, (2.5-)3-4.5 mm diam., with distinct, more 
or less prominent, fine reticulum, with slightly to 
rather strongly prominent, blunt to rather sharp me- 
dian keel which does not run out into a ventral pro- 
cessus or tubercle; ventral pore not sunken, some- 
times a bit elevated. 

Dist r . Sri Lanka throughout SE. Asia to China, 
S. Korea, Japan and Taiwan; in Malesia: N. Sumatra 
(Karo), Malay Peninsula, W. & Central Java, Philip- 
pines (Batan Is., Luzon, Mindanao). Rare in W. 
Malesia. Fig. 13. 

Ecol . Primary or secondary montane rain-forest, 
600-2500 m altitude, on loamy or volcanic soils, also 
on limestone if the climate is wet enough. At higher 
altitudes the subspecies is deciduous. In Malesia but- 
tresses are sometimes developed, up to 1.5 m high. 

Field notes . Bark dark to light grey, smooth, in 
old trees sometimes distantly shallowly fissured. In- 
ner bark soft, fibrous, with 'fingers' tapering out- 
wards into granular tissue, pale pinkish brown to dull 
red or redbrown, also said to be white, and turning 
salmon red on exposure. Wood light and soft, 
fibrous, easily split, white, with large pores and 
beautiful grain, with prominent rays, hcarlwood in 
older trees striped reddish and white. Leaflets be- 


Flora Malesiana 

[ser. I, vol. \0^ 

neath pale green, often glaucous. Fruits said to be 
reddish, green brown, or black when ripe. 

Vern. Sumatra: kabung sillang bulung, Batak 
lang.; Java: dangdur bulu, kawayang, ki sum, ki 
tiwu lalaki, S; Philippines: adope. adupong, 
aropong. bantinan, kamug, Ig., bae, If. 

Notes. Attention should be given to the relation 
between ssp. arnottiana and ssp. pendula in the 
Philippines (for the relation to ssp. macrophylla, see 
under that subspecies). In the mountains of Luzon 
both subspecies have been collected, ssp. pendula 
above 1800 m altitude and ssp. arnottiana from c. 
800-900 m up to c. 2400 m. Locally, e.g. on Mt San- 
to Thomas, they have been found together, but 
doubtless intermediate specimens are not observed. 
It is possible that in such localities these subspecies 
mutually behave as species; population studies in the 
field might yield more evidence with regard to this. 

The same problem arises in W. Malesia, where 55/?. 
arnottiana has been collected (rarely). In Sumatra 
and in Java its relation to ssp. ferruginea is in- 
teresting since there is an altitudinal zone of overlap 
between both, though ssp. ferruginea generally oc- 
curs lower than ssp. arnottiana. In Java the situation 
is as follows: ssp. ferruginea is by far the most com- 
mon of both, ssp. arnottiana having only been col- 
lected on a few mountains. Of these it is only G. Sa- 
lak and G. Gedeh where both subspecies have been 
found. Only of G. Gedeh more detailed ecological 
evidence is available: Koorders (FI. Tjibodas 2, 
1923, 157) stated that ssp. ferruginea occurs at c. 
1400 m altitude and that ssp. arnottiana ('M. glau- 
ca') occupies a zone between 1800 and 2400 m, being 
especially abundant at c. 2200 m. This does suggest 
the existence of ecological differentiation, but since 
ssp. arnottiana on other mountains also grows at 
lower altitudes, the situation remains unclear. 

b. spp. ridleyi (King) Beus. Blumea 19 (1971) 505. - 
Meliosma ridleyiKwG, J. As. Soc. Beng. 65, ii (1896) 
458; Ridley, J. Str. Br. Roy. As. Soc. n. 33 (1900) 
67; FI. Mai. Pen. 1 (1922) 516. - Meliosma elegans 
Ridley, J. Str. Br. Roy. As. Soc. n. 54 (1910) 40; FI. 
Mai. Pen. I (1922) 515. - Meliosma paucinervia 
Merr. Philip. J. Sc. 10(1915) Bot. 39; Enum. Philip. 
FI. PI. 2 (1923) 518. - Meliosma trichocarpa Merr. 
Pap. Mich. Ac. Sc. 24 (1938) 80, nom. illeg., non 
Hand.-Mazz. (1934). - Meliosma bartlettii Merr. 
& Perry, J. Am. Arb. 20 (1939) 356. - Meliosma 
confertiflora Merr. & Perry, I.e. 359. 

Shrub or tree, up to c. 20 m. Leaves 3-7-jugate; 
leaflets oblong to lanceolate, small to usually me- 
dium-sized, up to c. 20 by 6 cm, base acute, rarely 
rounded, usually entire, densely villous to glabrous, 
without domatia. Panicles erect, usually rather lax 
and slender; lower primary side-axes mostly subtend- 
ed by small leaves. Sepals and outer petals glabrous. 

Inner petals with fimbriate or ciiiolate tips, rarely 
glabrous. Ovary densely pubescent. Endocarps 
subglobose to very depressed and oblique, 4.5-9 
(- 10) mm diam., with vague to distinct, more or less 
prominent, rather wide reticulum, with slightly to 
strongly prominent, blunt to very sharp median keel 
which often at one end runs out into a minute ventral 
processus, the curving at the other end sometimes far 
drawn out into a blunt beak; ventral pore hardly to 
rather deeply sunken. 

Distr. Malesia: Central Sumatra, Malay Penin- 
sula, Borneo (Sarawak, Sabah, W. Kutai), Philip- 
pines (Mindoro). Fig. 13. 

Ecol . Primary and secondary rain-forest, both in 
mixed dipterocarp and in heath forest, on various 
soil types, from sea-level up to 1400 m altitude. 

Field notes. Bark mostly smooth, sometimes 
somewhat scaly or slightly fissured, grey to brown. 
Inner bark fibrous, pinkish to red or redbrown, 
turning brown after exposure. Young branches, 
inflorescence-axes, and leaf-rachises are sometimes 
(Singapore) covered with a dense layer of soft dark 
reddish brown hairs. Sepals sometimes said to be 
purple. Fruit often ± hairy ( 'trichocarpa'), once said 
to be bright purple. 

Vern . Sumatra: kaju rokkam, k. rube gala, k. si 
hasur, k. si (mardjuhut) (nij manuk, Asahan, mo- 
dang halimponan, Tapanuli. 

Notes . Ssp. ridleyi is rather variable when com- 
pared to the other subspecies of A/, pinnata, especial- 
ly in number and dentation of leaflets, in the degree 
of pubescence, and in shape and size of the endo- 
carps. In the Malay Peninsula, for instance, a form 
with few subglabrous and somewhat dentate leaflets 
has been found ('M. elegans'), as well as a beautiful, 
densely rufous-pubescent form with distinctly more 
and entire leaflets ('M. ridleyi'). It is not astonishing 
that such different plants have been described as 
separate species; only by studying material from 
Borneo it becomes clear that these extremes are con- 
nected by a range of transitions. Another form from 
Dallas (Kinabalu), which has rather condensed 
panicles, has been described as M. confertiflora. 
This again is merely a local form without any 
systematical significance, as is M. paucinervia, with 
very lax panicles, from Mindanao. Yet, in spite of 
this variation, it is obvious that ssp. ridleyi is a natu- 
ral unit, probably most closely related to the adjacent 
ssp. arnottiana from which it differs least of all 
subspecies, mainly in shape and size of the en- 
docarps, but also in some less important characters; 
an especially close resemblance has been observed 
between ssp. ridleyi and some deviating specimens 
from South Vietnam which have been tentatively in- 
cluded in ssp. arnottiana. Furthermore, the area of 
ssp. ridleyi borders on or somewhat overlaps the 
areas of ssp. ferruginea and macrophylla. The rela- 


Sabiaceae (van Beusekom & van de Water) 


tion between ssp. ridleyi and these subspecies has 
been discussed under ssp. macrophylla. 

Finally, it should be noted that the area of ssp. 
ridleyi fully overlaps that of M. sarawakensis; this is 
not accidental, since the latter probably is a 
derivative of ssp. ridleyi (see the note under M. 

c. ssp. ferruginea (Blume) Beus. Blumea 19 (1971) 
507. - Meliosma ferruginea Blume, Cat. (1823) 32, 
non SiEB. & Zucc. e.x Hook./. (1876), nee Kurz ex 
King (1896); Nees, Flora 8 (1825) 106; Hassk. Cat. 
Hort. Bog. (1844) 226; Blume, Rumphia 3 (1849) 
200; Walp. Ann. 2 (1852) 225; MiQ. Fl. Ind. Bat. 1, 
2 (1859) 616; Illustr. (1871) 74; K. & V. Bijdr. 9 
(1903) 121; KooRD. Exk. Fl. Java 2 (1912) 546; Atlas 
2 (1914) t. 375; Fl. Tjibodas 2 (1923) 157; Backer & 
Bakh./. Fl. Java 2 (1965) 145, p.p., exct. M. glauca 
et Jloribunda. - Millingtonia ferruginea Schult. & 
ScHULT. Syst. Veg. Mant. 3, add. 2 (1827) 250; 
DiETR. Syn. PI. 1 (1839) 103. 

Medium-sized to big tree, up to c. 42 m. Leaves 
2-6(-7)-jugate; leaflets elliptic to oblong, basal ones 
sometimes a bit ovate, upper ones sometimes ± 
obovate, usually rather large, up to 25(-38) by 
10(-18) cm, base rounded to truncate, sometimes 
acute, entire, firmly coriaceous, pubescent, rarely 
subglabrous, rarely with domatia. Panicles erect, 
spreading, lax to rather dense; lower primary side- 
axes usually subtended by small leaves. Sepals usual- 
ly densely pubescent, rarely on the outside sparsely 
so to subglabrous. Outer petals pubescent outside, 
rarely glabrous. Inner petals with fimbriate or 
ciliolate tips. Ovary partly or entirely but almost 
always distinctly and densely pubescent, very rarely 
nearly glabrous. Endocarps subglobose, often some- 
what depressed and oblique, 3.5-5.5(-8) mm diam., 
with rather vague to distinct, ± prominent reticulum, 
with usually very prominent, rather sharp median 
keel which does not run out into a ventral processus or 
tubercle; ventral pore not or not much sunken. 

Distr. Malesia: N. & Central Sumatra, through- 
out Java, and the Lesser Sunda Islands (Bali, Sum- 
bawa, Flores, Timor), locally common, especially in 
Java. Fig. 13. 

Ecol. Rain-forest, preferably on fertile, often 
volcanic soils, 250-1600 m altitude. 

Field notes. Bole cylindrical, straight, some- 
times crooked, at the base up to c. 2.5 m diam. Bark 
on the surface grey to brown, smooth, sometimes a 
bit peeling or shallowly fissured to (deeply) cracked, 
about 0.7-1.5 cm thick, easily detachable. Inner 
bark pale brown to brownrcd or orange, with 
streaks, also said to be dirty while and turning orange 
brown when exposed to the air as a result of the 
discolouring of the initially colourless watery exuda- 
tion. Wood soft, yellowish to pinkish white. Leaflets 

pale greyish to glaucous green beneath. Fruits 
brownred to black when ripe. 
Uses . Advocated for reafforestation purposes by 


Vern. Sumatra: sekapong, Takengon, sontang, 
Simelungun, sihubung, Kerinci; Java: ki tiwu, ki 
tjerme badak, S, gempong, gijubuk, gompong, J; 
Lesser Sunda Is.: gempong, sambuk, Bali, mladja, 
tanggo, tawu, Flores, Endeh lang., lohot, raok, 
Flores, kaju mangkok, W. Sumbawa. 

Note. Ssp. ferruginea is usually well recognizable 
by its outside pubescent sepals and petals. However, 
in N. Sumatra and the Lesser Sunda Islands 
specimens occur in which these characters are im- 
perfectly or not developed, and they may also lack 
the pubescence on the ovary and may have almost 
glabrous leaves. They are not easily identifiable and 
may be confused with A/, lanceolata var. lanceolata 
f. nervosa or with the closely related M. pinnata ssp. 
macrophylla and ssp. ridleyi. 

A. ssp. macrophylla (Merr.) Beus. Blumea 19(1971) 
510. - Meliosma macrophylla Merr. Philip. J. So. 
7 (1912) Bot. 294; Enum. Philip. Fl. PI. 2 (1923) 517. 

- Meliosma lanceolata var. obliqua {non Blume) 
KooRD. Minah. (1898) 408; Suppl. 2 (1922) 7, t. 55; 
ibid. 3 (1922) 28. - Meliosma wallichii (non Planch. 
ex Hook./.) Koord. Minah. (1898) 408. - Meliosma 
tongcalingii Elmer, Leafl. Philip. Bot. 8 (1915) 
2815. - Meliosma megalobotrys Mekr. Philip. J. So. 
11(1916) Bot. 16; Enum. Philip. Fl. PI. 2 (1923) 517. 

- Meliosma macrocarpa Elmer, Leafl. Philip. Bot. 
10 (1939) 3786, descr. angl. - Meliosma ferruginea 
(non Blume) Merr. & Perry, J. Arn. Arb. 20(1939) 
356. - Fig. 12. 

Medium-sized to large tree, up to c. 42 m. Leaves 
(3-)5-9-jugate; leaflets elliptic to oblong to ovate- 
oblong, medium-sized to rather large, up to c. 20 by 
9 cm, base rounded or obtuse to truncate, entire, 
rarely with a few teeth (Sulawesi), chartaceous to 
firmly coriaceous, very sparsely to densely pubes- 
cent, always without domatia. Panicles erect and 
spreading, lax and slender to rather dense; lower 
primary side-axes usually subtended by small leaves. 
Sepals and petals glabrous. Ovary glabrous, rarely 
with a few scattered hairs. Endocarps subglobose, 
sometimes more obovoid or depressed, more or less 
oblique, 3.5-5 mm diam., exceptionally 5-7.5 mm 
diam. (Sulawesi), with vague to distinct and promi- 
nent reticulum, with rather sharp and prominent me- 
dian keel which at one end mostly runs out into a 
small but distinct ventral processus or tubercle; ven- 
tral pore .somewhat sunken. 

Distr. Malesia: E. Borneo (E. Sandakan, Bcrao, 
W. & E. Kutai, Tandjung), Sulawesi (Minahasa, 
Malili), Moluccas (Halmahcra, Scram). Philippines 
(Luzon, Lcytc, Mindanao. Palawan), throughout 


Flora Malesiana 

[ser. I, vol. lO-* 

New Guinea (incl. New Britain). Fairly common in 
most parts of the area. Fig. 13. 

Ecol. Usually in primary, rarely in secondary 
rain-forest, at low to medium altitudes; in Borneo 
only collected below 100 m. in the other parts of the 
area also higher, up to c. 1 100-1200 m, in W. New 
Guinea once at 1800 m. in Borneo usually found in 
lowland dipterocarp forests. Generally reported to 
occur on clayish, loamy, or sandy clayish soils, also 
on red earth, on volcanic soil, and on loamsoil on 
limestone. It is rarely found in occasionally sub- 
merged areas. Once (New Guinea) said to occur on 
peaty soil, and there developing stiltroots. 

Field notes. Bole mostly straight, cylindrical, 
up to at least 1 m diam. at the base, usually develop- 
ing 1.5-2.5 m high buttresses, sometimes without 
buttresses, once observed stiltrooted. Bark grey to 
brown, or patchy brown-white-grey, smooth, some- 
times with shallow vertical cracks, not or little peel- 
ing, with vertical rows of lenticels. Inner bark c. 1 cm 
thick, soft, light brown or pink to brownred, inside 
paler, sometimes said to be streaked with cream, with 
some colourless sticky exudate (which is also said to 
be redbrown!); it is said to be rapidly darkening upon 
exposure or 'a bright orange-brown stain quickly ap- 
pears between bark and sapwood.' Wood very light 
and soft; sapwood white to pale pink or brown, when 
fresh with bright brown sap streaks, heartwood ab- 
sent or present, darker than the sapwood. The fruit 
is said to be brown to black. 

Vern. Borneo: surian, E. Kutai; Sulawesi: kaju- 
saul-rintek, Tooelooe lang., papako, Tontemboan 
lang., mumping, Tonsea lang., Hasan, Ratahan 
lang.; Moluccas: bais, Seram; Philippines: arocong, 
Ig., agosos, balilang-uak (a corruption of barilan ng 
uak). Tag., morau, S.L.Bis., mungapong, Bik., ma- 
gasorod. Bag.; New Guinea: sebotebuk, tubuk, 
Mooi, serajema, Manikiong, marwaskeipi, Japen, 
bagare, Kapauku lang., biedewon, iediewat, Muju, 
morrotuno, waito, Wapi, frikipa, Orne lang., 
tapuha, Managalase, kufi, Kutubu, uliga, Madang, 
kombowase, Waskuk, wagebi, Wagu. 

Notes. Ssp. macrophylla is the most common 
and widespread of the East Malesian subspecies 
group, characterized by a glabrous ovary by which it 
is readily distinguished from the West Malesian 
subspecies. Within its large area a few other sub- 
species occur, viz. ssp. pendula, sylvatica, and 
humilis, which have much more limited areas and 
probably represent offsplits from it. These three 
subspecies are ecologically well isolated from ssp. 

In Borneo the area oissp. macrophylla is, as far as 
can be judged from the available evidence, sharply 
delimited against that of the West Malesian ssp. 
ridleyi, which, moreover, appears to prefer a higher 
altitudinal zone (only in Borneo ssp. macrophylla 

seems to be restricted to lowland forests below c. 100 
m altitude!). To the SW the area o^ ssp. macrophylla 
borders on that oi ssp. ferruginea which inhabits the 
Lesser Sunda Islands. The latter two subspecies are 
huge trees, very similar in general habit, and some- 
times they have been confused. Nevertheless, they 
are usually well distinct, mainly by flower characters, 
though in both subspecies there is a tendency to lose 
some of these characters. 

In the Philippines ssp. macrophylla is sympatric 
with ssp. arnottiana, but they prefer different 
altitudinal zones, the first being a lowland subspecies 
not exceeding c. 900 m altitude, the latter being a 
montane subspecies occurring from c. 800 up to c. 
2400 m (once recorded from c. 600 m). 

e.spp. pendula (Merr.) Beus. Blumea 19(1971)512. 

- Meliosma pendula Merr. Publ. Govt. Lab. 
Philip. 29 (1905) 25; Enum. Philip. Fl. PI. 2 (1923) 
518. - Meliosma reticulata Merr. Philip. J. Sc. 5 
(1910) Bot. 195; Enum. Philip. Fl. PI. 2 (1923) 518. 

- Meliosma macgregorii Merr. Philip. J. Sc. 10 
(1915) Bot. 37; Enum. Philip. Fl. PI. 2 (1923) 517. 

Small to medium-sized tree, up to c. 20 m. Leaves 
(3-)4-6-jugate; leaflets elliptic to oblong, rarely lan- 
ceolate, up to 18(-20) by 7(-ll) cm, the lower ones 
at the base nearly always rounded or (sub)truncate to 
obtuse, the upper ones more or less acute, nearly 
always distinctly dentate, villous-pubescent, ± gla- 
brescent, without domatia. Panicles erect and 
spreading to somewhat pendulous and rather flaccid, 
slender and rather lax; lower primary side-axes most- 
ly subtended by small leaves. Sepals and petals 
glabrous. Ovary glabrous. Endocarps subglobose, 
slightly oblique, 6-7 mm diam., with rather vague, 
slightly elevated reticulum, with hardly to moderate- 
ly prominent, blunt median keel, the latter not run- 
ning out into a distinct ventral processus or at most 
into a very minute tubercle; ventral pore hardly or 
not sunken. 

Distr. Malesia: Philippines (Luzon: Mountain 
Prov.). Fig. 13. 

Ecol. Montane rain-forest, 1800-2500 m alti- 
tude. In mossy forest, in ravines as well as on expos- 
ed ridges. 

Field notes. Bark thick, checked. Wood soft, 
said to be soon assuming an orange-brown colour. 

Uses. The leaves are once said to be used for 
smoking by the Igorots. 

Vern. Anitap, Ig. 

Note. Ssp. pendula replaces the lowland and 
lower hill ssp. macrophylla at high elevations (in this 
respect being comparable to ssp. humilis from New 

f.ssp. sylvatica (Elmer) Beus. Blumea 19(1971)513. 

- Meliosma sylvatica Elmer, Leafl. Philip. Bot. 


Sabiaceae (van Beusekom & van de Water) 


2 (1908) 492; Merr. Enum. Philip. Fl. PI. 2 (1923) 
518. - Meliosma acuminatissima Merr. Philip. J. 
Sc. 10 (1915) Bot. 36; Enum. Philip. Fl. PI. 2 (1923) 
517. - Meliosma brachyboirys Merr. Philip. J. Sc. 
12 (1917) Bot. 275; Enum. Philip. Fl. PI. 2 (1923) 

Slender shrub or treelet, up to c. 5 m. Leaves 
(3-)4-6(-7)-jugate; leaflets elliptic to usually ob- 
long or lanceolate, usually medium-sized, up to c. 18 
by 6 cm, acute at the base, sparsely to rather closely, 
always distinctly dentate, very sparsely to moderately 
pubescent, without domatia. Panicles erect, spread- 
ing, usually slender and rather la,\; primary side-axes 
(mostly?) not subtended by small leaves. Sepals and 
petals glabrous. Ovary glabrous. Endocarp subglo- 
bose, often somewhat ellipsoid, more or less oblique, 
5-6 mm diam., with distinct, more or less prominent 
reticulum, with rather sharp and prominent median 
keel which at one end runs out into a small but 
distmct ventral processus; ventral pore somewhat 

Distr. Malesia: Sulawesi (Minahasa, Latimod- 
jong Mis), Philippines (Luzon, Negros). Fig. 13. 

Ecol . Lowland rain-forest, usually not above 750 
m altitude, growing in the shrub layer. 

Field notes. Slender, suberect or bent shrub or 
treelet of a sparsely branched habit. Bark smooth, 
grey and brown mottled. Wood white, soft, easily 
breakable. Leaves once said to be light bluish green 

Note. Ssp. sylvatica is closely related to ssp. 
macrophylla. The most striking difference between 
them is found in their physiognomy, the former be- 
ing a small undergrowth treelet, the latter a large 
forest tree. The main systematical differences are 
found in the dentation and the shape of the base of 
the leaflets. These fit nicely in the spectre of 
character combinations present in the subspecies of 
M. pinnata, and it seems justified to consider ssp. 
sylvatica a subspecies of that species, instead of a 
separate species. 

R. spp. humilis (Merr. & Perry) Beus. Blumea 19 
(1971) 514. - Meliosma humilis Merr. & Perry, J. 
Arn. Arb.20(1939)358;/*/Vy. 22 (1941) 263, //j 065. 
- Meliosma schlechteri Merr. & Perry, J. Arn. 
Arb. 22(1941) 262. 

Small to medium-sized tree, up to c. 20 m. Leaves 
3-5 jugate; leaflets elliptic to oblong, rarely short- 
lanceolate, usually ralhcr small, up to c. 15(~24) by 
6<-9) cm, the base acute, lower leaflets sometimes 
rounded at the base, sparsely to rather closely den- 
tate, beneath subglabrous to ralhcr densely pubes- 
cent, sometimes densely villous-tomcntose, always 
with more or less distinct domatia in the axils of the 
nerves beneath (obscure in densely lomentose 
leaflets). Panicles erect, spreading, mostly rather lax 

and with slender axes; lower primary side-axes often 
subtended by small leaves. Sepals and petals gla- 
brous. Ovary glabrous. Endocarps subglobose, 
somewhat depressed and rather oblique, 5.5-7.5 
mm diam., with more or less vague, slightly elevated 
reticulum, with very prominent, rather sharp median 
keel which at one end runs out into a small but 
distinct ventral processus; ventral pore somewhat 

Distr. Malesia: Papua New Guinea (Highlands 
Provinces, Madang, Morobe, Milne Bay). Common. 
Fig. 13. 

Ecol. Montane rain-forests, 1000-30(X) m 
altitude. Observed as an understorey tree in dense 
Castanopsis-Nothofagus forest, on ridges as well as 
on streambanks, but also often reported from several 
kinds of disturbed forest, such as bamboo regrowth, 
old garden land, transition between coniferous forest 
and treefern grassland, and even from open grass- 
land. Once reported from limestone ridge. 

Field notes. Bark greybrown, smooth, with big 
lenticels. Inner bark straw-coloured to pink, red, or 
reddish brown (due to discolouring, as in other sub- 
species?), exuding 'resin'. Wood white to light 
brown, with conspicuous rays and clear growth 
rings, said to be of moderate weight and hardness. 
Petioles, peduncle, and pedicels purplish to red- 
brown; buds reddish. Fruits dark red to black when 

Uses. Once said to be used as housing timber, 
free from borers. 

Vern. Mansalong, Finschhafen, kokopong, 
Nako lang., E. Madang Prov., kass, Maring lang., 
mappam, Enga lang., W. Highl. Prov. 

Notes. Ssp. humilis is closely allied to ssp. ma- 
crophylla, mainly differing by its dentate leaflets 
with domatia. In Papua New Guinea it replaces ssp. 
macrophylla mainly found below 1000 m altitude (cf. 
ssp. pendula from Luzon). It is remarkable that ssp. 
humilis has as yet been collected, even rather abun- 
dantly, only in Papua New Guinea and not in W. 
New Guinea. 

7. Meliosma sarawakensis Ridley, Kew Bull. (1933) 
193; Merr. & Perry, J. Arn. Arb. 20 (1939) 359. - 
Meliosma grandifolia Lecomte, Bull. Soc. Bot. Fr. 
54(1909)676, now. ///e/?.. non Urban (1895); Merr. 
Enum. Born. (1921) 362; van Beusekom. Blumea 19 
( 1 97 1 ) 5 1 5 . - Meliosma confusa var. laxior Baker /. 
in Rendle, J. Bot. 62 (1924) Suppl. 30. - Meliosma 
lattfolia RitUEY. Kew Bull. (1933) 193; Merr. & 
Perry, J. Arn. Arb. 20 (1939) 359. m obs. 

Evergreen, small tree, up to c. 10 m. Leaves 
2-3(-4)-jugate; rachis terete, 12-30 cm, including 
the 6-15 cm long petiole, up to c. 5 mm diam.. 
densely short-tomcntosc. later ± glabresccni; leaf- 
lets usually elliptic to oblong, the lower ones often 


Flora Malesiana 

[ser. I, vol. 10^ 

more or less ovate to ovate-oblong, the upper ones 
often more or less obovate to obovate-oblong, 
sometimes ± asymmetrical, (2-)5-22 by (1.5-) 
3-12 cm, mostly distinctly increasing in size towards 
the top of the leaf, base acute to rounded, apex more 
or less acuminate, sometimes subacute or cuspidate, 
with entire to remotely spinously dentate margin, 
chartaceous, moderately to rather densely pubescent 
especially beneath and on midrib and nerves, often 
partly giabrescent when older, never with domatia; 
midrib more or less impressed above; nerves 6-12 
pairs, ascending, usually looped; venation distinct, 
reticulate; petiolules up to c. 1.5(-3) cm, terminal 
one usually longest, tomentose. Panicles terminal, 
usually more or less pendulous, flaccid, lax, narrow- 
ly pyramidal, (20-)25-55 cm, not profusely 
branched up to the 2nd or 3rd order, branches 
spreading, ± flaccid, usually slender, densely 
tomentose, bearing numerous flowers crowded in 
dense spikes; primary side-axes few to rather many, 
up to c. 25(-35) cm, the lower ones usually subtend- 
ed by reduced leaves; bracts ovate to usually narrow- 
ly triangular or linear-lanceolate, up to c. 4 mm, 
densely pubescent. Pedicels (almost) absent. Mature 
buds c. 2 mm diam. Sepals 5 (4), ovate to ovate- 
lanceolate, the 3 or 4 inner ones 1-1.5 mm, the outer 
1 or 2 usually much smaller, often minute, densely 
pubescent on the outside, with entire margin. Outer 
petals glabrous. Inner petals about halfway or some- 
what less bifid, 0.5-0.7 mm, glabrous, sometimes 
with a minute central lobule. Filaments c. 1 mm. 
Ovary 0.5-0.7 mm, densely pubescent. Fruit 
(sub)globose, when ripe 0.7-1 cm diam.; endocarp 
depressed-globose, applanate at the ventral side, 
strongly oblique, 6-7(-8) mm diam., with usually 
distinct, more or less sharply prominent reticulum; 
median keel sharp and very prominent, not at one 
end running out into a ventral processus or tubercle, 
at the other end rather far curving outwards; ventral 
pore rather sunken. 

Distr . Malesia: Sumatra (Asahan to Palembang) 
and NW. Borneo (Sarawak, around Kuching and 
Pontianak; common). Fig. 13. 

Ecol. Lowland rain-forest, up to c. 800 m 

Field notes. The sepals are redbrown to purple. 

Vern. Sumatra: kaju rube boras, Asahan; Bor- 
neo: bulu manok, Iban name, Kuching. 

Note. The closest affinity of M. sarawakensis is 
doubtless with M. pinnata ssp. ridleyi to which it is 
very similar in all characters (they even share the red 
sepals). Only after some hesitation M. sarawakensis 
is maintained as a separate species and not made a 
subspecies of A/, pinnata. It would fit rather well into 
that species but is distinguished from it by a wider 
range of characters. The most important of its 
characters are the 2- or 3-jugate leaves and the dense- 

ly pubescent sepals. These characters indeed are also 
found in M. pinnata ssp. ferruginea, but this subspe- 
cies is quite different from M. sarawakensis in 
various other aspects. An additional argument to the 
specific status of M. sarawakensis is found in the fact 
that it is found together with M. pinnata ssp. ridleyi 
in the same area and at the same altitudes in Sarawak 
(near Kuching) and in Sumatra (Asahan), without 
any sign of hybridization. 

8. Meliosma rufo-pilosa Hend. Card. Bull. Str. 
Settl. 7 (1933) 96, t. 18; Merr. & Perry, J. Arn. 
Arb. 20 (1939) 360; van Beusekom, Blumea 19 
(1971) 517. 

Evergreen rather large tree up to c. 30 m. Flower- 
ing twigs terete, 5- 10 mm diam., stout, abruptly ter- 
minating in a tuft of leaves and inflorescences, gla- 
brous, often with many large conspicuous leaf-scars. 
Leaves (6-)7-9-jugate; rachis terete, (13-)25-50 
(-65) cm, including the (3.5-)6- 16 cm long petiole, 
pubescent, not swollen at the base, hardly or not len- 
ticellate; leaflets elliptic to oblong, sometimes ovate 
(-oblong), 3- 15 by 2-6 cm, base obtuse to truncate, 
apex acuminate, entire, glabrous or ± puberulous 
on nerves above, (sub)glabrous beneath, pubescent 
on nerves, always without domatia; midrib im- 
pressed above; nerves 7-18 pairs, ascending, looped; 
venation fine, very distinct, reticulate; petiolules 1-6 
mm, densely pubescent. Panicles terminal, one or a 
few crowded together at the end of a twig, erect, 
rather dense to lax, pyramidal, 30-50 cm, including 
the 0-20 cm long peduncle, profusely branched up to 
the 4th order, branches spreading, ± flaccid, densely 
pubescent, bearing numerous solitary flowers; 
primary side-axes well-spaced, c. 8-15, up to c. 30 
cm, not lenticellate, the lower ones never subtended 
by small or reduced leaves; bracts narrowly triangu- 
lar to lanceolate, up to c. 4 mm, densely pubescent. 
Pedicels 1-3 mm, densely pubescent. Sepals (3) 4, 
ovate, (sub)equal, c. 1.5-2 mm, the outer one often 
much smaller, rarely minute, usually lowered on the 
pedicel, glabrous or somewhat pubescent outside; 
margin flimsy, more or less ciliolate, entire or some- 
times with some coarse irregular teeth. Outer petals 
c. 1.5 by 1.5-2.5 mm. Inner petals ligular, usually 
somewhat widened towards the top, 0.7-1 mm; top 
entire or with a shallow incision, blunt, minutely 
ciliolate. Filaments 0.7-1 mm. Ovary 0.5-0.7 mm, 
glabrous. Fruit globose, when ripe 1.5-2 cm diam., 
with moderately thick, fleshy mesocarp; endocarp 
semiglobose, broad-ovate to subcordate at ventral 
view, 11-13 mm long and wide, 7-8 mm high, with 
relatively thin wall, with slightly lumpy surface, 
especially lumpy and somewhat furrowed at the ven- 
tral curving of the wall; median keel faint, hardly 
elevated but at one end drawn out into a con- 
spicuous, laterally flattened, downwards-curved. 

1989] Sabiaceae (van Beusekom & van de Water) 715 

blunt beak; ventral side rather deeply concave with a crown, once reported with c. 2 m high buttresses, 

smooth, wide-ovate to suborbicular central part Bark smooth, grey to brown, with lenticels in vertical 

from the centre of which protrudes the ± conical rows {'scarred', 'dippled'). Inner bark soft, fibrous, 

hilum of the seed. orange to reddish outside, pale fawn to white to- 

Distr. Malesia: Malay Peninsula (Pahang), wards the cambium. Sapwood pale brown. Twigs 

Borneo (Sarawak, Sabah, Kinabalu complex). Fig. pale brown, rough, lenticellate, with darker leaf- 

1 1 . scars. Leaves pale green. Fruit yellow to orange when 

Ecol. Montane rain-forest, 1350-1700 m alti- ripe, 

tude. Vern. Malay Peninsula: sengkuang, Genting 

Field notes. Large tree with deep, rounded Highlands. 

Excluded and dubious 

Meliosma celebica Warb. ^.v Dihm, Beih. Bot. Centralbl. 21, 1 (1907) 125, nomen. - I have not seen the 
type specimen (Warburg 15416, Sulawesi, Bojong), which probably got lost during World War \\. 

Meliosma laurina Bllme, Rumphia 3 (1849) 198; Walp. Ann. 2 (1852) 224; Miq. F1. Ind. Bat. 1, 2 (1859) 
614; lllustr. (1871) 73; Merr. Enum. Born. (1921) 363; Hall./. Beih. Bot. Centralbl. 39, 2 (1921) 161; 
KosTER.M. Bibl. Laur. 1 (1964) 951. - As was noted by Hallier/., I.e., the type specimens (S. Muller^.a;., 
Borneo, G. Sakumbang) consist of a mixture, viz. inflorescences of M. sumatrana and leaves of Cryptocarya 
reticulata Blume (Lauraceae). 

Meliosma petiolaris Miq. Sum. (1861) 519, 203; lllustr. (1871) 73, in obs. - This species was later referred 
by MiQUEL himself to Xylosma leprosipes Clos which is now known as Bennettiodendron leprosipes (Clos) 
.Merr. (Flacourtiaceae). 

Meliosma timorensis Blume ex Blenk, Flora 67 (1884) 370, nomen. - This name was cited in an enumera- 
tion oi Meliosma species having leaves with pellucid dots; it was probably copied from a label on a sheet. The 
specimen could not be traced. 

Pimela angustifolia BivMt, Mus. Bot. Lugd.-Bat. 1 (1850)226. - Canariopsis angustifolia Blvme e.x Miq. 
Fi. Ind. Bat. 1, 2 (1859) 653. - Canarium angustifolium Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 117; 
H.J. Lam, Bull. Jard. Bot. Btzg III, 12(1932) 179. t. 11 f. l\d,subC. rigidum Zipp.; Leenh. F1. Males. 1, 
5 (1956)296. 

The material under this name was excluded from the Burseraceae by Leenhouts, I.e., and tentatively 
assigned to Meliosma. This may be correct, and it should then be placed close to M. laneeolata and M. hirsuta. 
At first sight it is very similar to the latter species, but there are important differences in nervation and 
pubescence. If it belongs to Meliosma it would certainly be a new species, but I refrain from including it 
because I am not sure about its identity. Unfortunately, the specimens consist of young leaves only, with many 
characteristic narrow leafiets, but in absence of woody parts it cannot be identified with certainty. Moreover, 
on the original labels (Zippel^./;.) the place of origin is mentioned as being 'Nova Guinea', in Blume's hand- 
writing. However, this would not fit in the distribution pattern of Meliosma, since species of this kind only 
occur in western Malesia; if New Guinea indeed is the correct locality, 'Canarium angustifolium' can hardly 
belong to Meliosma. Its identity will probably remain uncertain until more satisfactory material has been 


C. G.G.J. VAN SteenisI, W.J.J.O. de Wilde, c.s. 

As was done in the preceding volumes, it seemed useful to correct some errors which have crept into the text 
of volumes 4- 1 as well as to add some additional data, new records and references to new species which came 
to our knowledge and are worth recording. 

Volume and page number are separated by a colon. Page numbers provided with either a ot b denote the 
left and right columns of a page respectively. 


9: 47b Osmoxylon sessiliflorum (Laut.) Philip- 


Add to Distr.: Moluccas (Halmaheira), 2 


Add to Vern.: saha-saha, Ternate lang., 

tele, Sahu lang. 


8: 142b Dolichandrone spathacea (L.f.) K. Sch. 
Add to Distr.: A.G. Wells, in H.J. Teas 
(ed.). Biology and ecology of mangroves 
(1983) 61, map, records this species for the 
first time from Australia: the northern tip 
of Cape York Peninsula (2 localities). 


7: 93 Cardiopteris [Wall, ex] Royle. 

Add to Distr.: The genus extends to the 

Solomon Islands (Bougainville) and N. 

Queensland; cf. Bailey, Queensl. Fl. 

(1899) 251; Compreh. Cat. Queensl. Pi. 
(1912) 93, f. 76. 


6: 257b Glyptopetalum loheri Merr. 

Distr.: Philippines: add Palawan (Rids- 

DALE 683). 
6: 266a Lophopetalum beccarianum Pierre. 

Add to Distr.: Central Sumatra (Djambi: 

Vreeken-Buys 69). 
6: 291 b Perrottetia alpestris (Bl.) Loes. ssp. philip- 

pinensis (Vidal) Ding Hou. 

Add to Distr. (and map): Lesser Sunda 

Islands (Flores: Loeters 650, 1796). 


4: 434b Jacquemontia brown iana Ooststr. 

Add the synonym: J. pannosa (R. Br.) 
Mabberley, Bot. Macar. 6 (1980, '1978') 


7: 653a Cyperus diaphanus Schrader ex R. & S. 

var. latespicatus (Boeck.) Kern. 

Add to Distr.: Lesser Sunda Islands 

(Flores: Schmutz 5767). 
9: 173a Carex bilateralis Hayata. 

Add to Distr.: SE. to E. Asia. 


9: 436b Hopea gracilis Miq. (under Excluded). 
It was unfortunately overlooked that the 
type at L (W. Central Sumatra, Padang, 
Teijsmann HB 424) was already in 1968 
referred by Ashton to Meiogyne and later 
in the same year identified by F.H. Hilde- 
BRAND as Meiogyne virgata (Bl.) Miq. {An- 


6: 856b Vaccinium angulatum J.J.S. 1914, non 
(Griff.) Theobald, 1883 = Vaccinium 
commutatum Mabberley & Sleumer, 
Taxon 34 (1985) 155, nom. nov. 


5: 24b Hydnocarpus nana King. 

Add to Distr.: NE. Sumatra (Besitang R., 

Sikundur For. Res., de Wilde & de Wilde- 

Duyfjes 19540). 
5: 35a Scaphocalyx spathacea Ridl. 

Add the synonym: S. parviflora RrDL. etc. 
5: 35b Reduce Scaphocalyx parviflora Ridl. to S. 

spathacea Ridl. 
5: 56a Homalium dasyanthum. 

Change authorship: (Turcz.) Theob. in 

Mason, Burma ed. Theob. 2 (1883) 451; 

Warb. etc. (see Mabberley, Taxon 34, 

1985, 155). 
5: 98b Reduce Casearia pallida Craib. to C 

flavovirens Bl. 
5: l(X)b Casearia flavovirens ^i.. 

Add the synonym: C. pallida Craib, etc. 



Addenda, corrigenda et emendanda 


5; 113a 

Add to Distr.: Thailand; in Malesia: Suma- 
tra, Java, Lesser Sunda Islands (Bali). 


Haemodorum corymbosum Vahl should 
be called Haemodorum coccineum R.Br., 
according to T.D. Macfarlane, Perth, 


8: 83 The correct name for Gladiolus natalensis 
(Ecklon) Reinw. ex Hook, should be 
Gladiolus daienii Geel, Sen. Bot. fasc. 28 
(1829). See Milliard & Burtt, Notes Roy. 
Bot. Card. Edinb. 37 (1979) 297. 



5: 366 Maingaya malayana Oliver. 

A marvellous colour photograph of this 
very rare species (and genus), endemic in 
Malaya and Penang, was given by Francis 
Nc in Nature Malaysiana 7, no 4 (1982) 8; 
the species is thought to be ± extinct and is 
now grown at the Forest Research Institute, 


5: 390b. Blyxa aiiheriii Rich. 

391 Add: var. echinosperma (Clarke) Cook & 

LiJOND, Aquat. Bot. 15 (1983) 14. 

Add the synonym: B. echinosperma 

(Clarke) Hook./, elc. 
5: 392a Blyxa leiosperma Koidz. 

Reduced to B. japonica (Miq.) Maxim, ex 

AscHERS. & Gl'rke var. japonica. 
5: 392b Blyxa aliernifolia (Miq.) Hartog. 

Reduced to B. japonica (Miq.) Maxim, ex 

Aschers. & GuRKE var. alternifolia (Miq.) 

Cook & Luond, Aquat. Bot. 15 (1983) 25. 
5: 393a fl/vx^crvopoAj/co (Miq.) Maxim, ex Aschers. 

& Gurke. 

Add: var. japonica and var. aliernifolia 

(Miq.) Cook. & Luond, Aquat. Bot. 15 

(1983) 22 and 25 respectively. 

Add the synonyms: B. leiosperma Koidz. 

and B. alternifolia (Miq.) Hartog. 


8: 18b Replace Hypericum uralum Bif h.-Ham. 
ex D.Don by Hypericum henryi Lev. & 
Vaniot ssp. hancockii Robson, Bull. Br. 
Mus. Nat. Hist. Bot. 12(1985)261, map 22. 
Distr. of var.: Continental SE. Asia (S. 
China, Vietnam, Burma, Thailand). 


7: 6a Cilronelta suaveolens (Hi.) Howard. 

Add to Distr.: Lesser Sunda Islands 
(Flores: Schmutz 5819). 

10: 7a The correct name for Olax scandens Roxb. 
should be Olax psittacorum (Willd.) 
Vahl; see Almeida, J. Bomb. Nat. Hist. 
Soc. 81 (1985) 742. 

10: 9b E.\cluded, add: 

Olax baliculin Blanco, F1. Filip. ed. 2 
(1854) 589; ed. 3 (1877) 38; Kosterm. Bibl. 
Laur. (1964) 1153 = Litsea baliculin 
(Blanco) Kosterm. Bull. Bot. Surv. India 
10(1968) 268 (Lauraceae). 

Merrill (Enum. Philip. Fl. PI. 2, 1923, 
195) recognized it as a Litsea and said that 
it might prove to belong to L. leytensis 
Merr. Since no material is known to exist, 
the matter will remain speculative. 


7: 416a Passiflora aurantia Forst. /. 

Add to Distr.: W. Central Celebes, near 
Palu, at 200 m, Meijer 10172. 


5: 356b Pittosporum moluccanum (Lamk) Miq. 

Add the synonym: Coffea angustifolia 
RoxB.;c/. FoRMAN,Kew Bull. 38(1983)64. 


6: 66b Cladopus nymani H. Moll. 

Add to Distr.: Central Borneo, en route 
from Sinar Baru to Ryan Ruwan, N of 
Long Bawan, Krayan, 1 1 5° 45 ' E, 4° 5 ' N, 
on rock in a rapid stream (Pa Raya), sub- 
merged and sterile, 1150 m alt., M. 
Okamoto C.S., s.n. (L, Osaka). The 
material matches the figure on p. 66 of Fl. 
Males. I, 4 almost exactly. 


5: 461b Bruguiera gymnorrhiza (L.) Saviony. 

Distr. : It is worthy of record that according 
to an internal report by A.N. Gillkson 
(C'SIRO, Canberra) this mangrove species 
is found on the lower limestone terrace on 



Flora Malesiana 

[ser. I, vol. 10"^ 

Christmas I. (Indian Ocean), in a healthy 
and self-maintaining community near the 
treshwaler Hosnie's Springs al 40 m alt., 
and not tar from it a stand of a remnant ht- 
toral forest of Heriliera littoralis Dryand. 
in Ait., another mangrove species. 
5 : 471a Ceriops decandra. 

The authorship must be (Griff.) 
Theobald; see Taxon 34 (1985) 154. 


6: 218b Ailanthus inlegrifolia LAMk. 

Add to Distr. (and map): N. Queensland 
(several localities) (L. Pedley, in litt.). 


4: 233 The correct name for Sparganium simplex 
HuDS./. simplex as used by Backer should 

Sparganium fallax Graebn. in E. & P. Pfl. 
R. 2 (1900) 15, t. 3H; Cook & Nicholls, 
Bot. Helv. 96 (1986) 253; ibid. 97 (1987) 3, 
t. 13a, 14a, map 14. - S. simplex auct. non 
HuDS./. simplex: Backer, FI. Males. I, 4 
(1951) 233, fig. 

Basal leaves usually exceeding the stem, 
(4-)5-10(-15) mm wide. Lowermost bract 
20-35(-60) cm long, 1 -several times as 
long as the inflorescence. Female heads 
3-4(-6), usually supra-axillary, sometimes 
extending beyond the next internode; ped- 
uncle up to 3 cm long. Male heads 5-8 or 
more. Female flowers: perianth segments at 
least connate below, in fruit more than 0.5 
times as long as the fruit; pedicel 1-3 mm 

long. Male flowers: filaments up to 6 or 
more mm long; anthers (0.8-)l -2(-2.2) 
mm long. Fruits fusiform, sometimes con- 
stricted around the middle, 5-6 mm long, 
light brown, dull, tapering below into a 
1.5-3 mm long pedicel. Endocarp c. 2 mm 

Distr. E. India to Japan; in Malesia: 
Sumatra, Papua New Guinea (up to 2000 m 

Sparganium subglobosum Morong, Bull. 
Torrey Bot. Club 15 (1888) 81, t. 79, f. 1; 
Cook & Nicholls, Bot. Helv. 97 (1987) 4, 
t. 13b, 14b, map 15. 

Basal leaves usually exceeding the in- 
florescence, (l-)2-4(-9) mm wide. Low- 
ermost bract up to 15 cm long, 0.5-1 (rare- 
ly more) times as long as the inflorescence. 
Female heads solitary, axillary or on lateral 
branches, occasionally one head supra- 
axillary, sessile, rarely with an up to 5 mm 
long peduncle. Male heads solitary. Female 
flowers: perianth segments free, in fruit 
0.3-0.5 times as long as the fruit; pedicel 
O-I mm long. Male flowers: filaments 
2.5-3. 2(-3. 8) mm long, anthers 0.5-0.9 
(-1) mm long. Fruits obovoid to almost 
globose, yellowish to pale brown, shiny, 
subsessile or with an up to 1 mm long, not 
persistent pedicel. Endocarp 1.6-1.8 mm 

Distr. India (once, in Khasia), Yunnan, 
Vietnam to Manchuria and to Okinawa, 
Australia, New Zealand; in Malesia: Papua 
New Guinea. 

Note. The identity of a collection from 
Arfak is not resolved by Cook & Nicholls. 


8: 217 Symplocos J acq. subg. Hopea Clarke. 
Add in key couplet la as follows: 

1. Leaves (pseudo-)verticillate. 

la. Leaves obovate 55. S. verticillifolia 

la. Leaves elliptic. 

2. Upper side of leaves hairy. Twigs tomentose 30. S. herzogii 

2. Upper side of leaves glabrous. Twigs appressedly to patently short-hairy S. rayae 

1 . Leaves not verticillate, etc. 

8: 218 Add couplet 32a as follows: 

32. Leaves longer than 5 cm (mean length). 

32a. Leaves 14-36 by 5-11 cm. Twigs tomentose S. riangensis 

32a. Leaves 3- 13(- 18) by 1.5-4.5(-6)cm. If leaves longer than 13 cm or broader than 4.5 cm then 

twigs not tomentose. 
33. Flowers etc. 

1989] Addenda, corrigenda et emendanda 719 

8: 227 Add couplets 5a, 5b, 15a, and 16b as follows: 
4. Twigs hairy. 
5. Leaves distichous. 

5a. Leaves 14-36 cm long S. riangensis 

5a. Leaves at most 12 cm long 33. S. laeteviridis 

5. Leaves spirally arranged or pseudo-verticillate. 
5b. Leaves pseudo-verticillate or at least 3-5 close together at the end of the flushes . S. rayae 
5b. Leaves spirally arranged. 
6. Leaves etc. 

15. Leaves longer than 5 cm. 

15a. Inflorescence a fascicle S. iliaspaiensis 

15a. Inflorescence a (short) spike, raceme, or panicle. 

16a. Inflorescence a (compound) spike S. cochinchinensis ssp. laurina 

16a. Inflorescence a short raceme or a panicle 42 (3). S. ophirensis ssp. cumingiana 

14. Bracts eic. 

8: 230 Add couplet 25a as follows: 

25. Inflorescence a raceme but pedicels sometimes very short. Bracts caducous. 

25a. Leaves 1 5-22 cm long 36. S. maliliensis 

25a. Leaves 3-6 cm long S. ambangensis 

25. Inflorescence etc. 

8: 235 Add the couplets 10a, 22a, and 22b as follows: 

10. Inflorescence not a spike. 

10a. Inflorescence a fascicle S. iliaspaiensis 

10a. Inflorescence a raceme or panicle 42. S. ophirensis 

10. Inflorescence etc. 

22. Leaves distichous. 

22a. Leaves 14-36 cm long S. riangensis 

22a. Leaves at most 12 cm long. 
23. Underside etc. 
22. Leaves spirally arranged or pseudo-verticillate. 
22b. Leaves pseudo-verticillate or at least 3-5 close together at the end of the flushes . S. rayae 
22b. Leaves spirally arranged. 
29. Leaves eic. 

8: 237 Add couplet 17a in between couplet 17: 

17a. Leaves 15-22 cm long 36. maliliensis 

17a. Leaves 3-6 cm long S. ambangensis 

or next to S. cochinchinensis as couplet 20a: 

20. Stone different. 

20a. Leaves more than 6 cm long 16. S. cochinchinensis 

20a. Leaves shorter than 6 cm S. ambangensis 

f&mWy Symplocaceae Symplocos costalifrucia Nom. Blumea 31 (1986) 

277, f. 1. 
Add the following species: Dislr. Malesia: Borneo (Sarawak, Brunei, 

S>mplocos rayae Nrxn. Blumea .30 (1984) 73. Symplocos iliaspaiensis Noot. Blumea 30(1984) 279. 

D.str, Malesia: Borneo (Kalimantan Tcngah. Di si r . Mfl/rs/a: Borneo (Sarawak and possibly E. 

Bukit Raya, only known from the type collection). L-,.i;.,r....,i..r,» 

Null in till lull /. 

s\mpl<Kos riangensis N^kh. Blumea 30 (1984) 74. Symphuos a 

Disir. Malesia: Borneo (Kalimantan Tcngah, 263,1. 1. 
I', it 11 Raya). Oislr. Malesia: North Sulawesi (Mcnado, Poso). 


Flora Malesiana 

[ser. I, vol. 10^ 


10: 109 According to Ms. T. Rubsamen (Bochum, 
West Germany; in litt.) the following 
amendments should be made: 

10: 110 Family distribution map: extend the central 
area to the west to include part of East 
Africa; see Kew Bull. 36 (1982) 733. 
Sciaphila Blume. 

Add/change description: Anthers some- 
times 4-celled (America). Endosperm pres- 
ent; embryo small, orthotropous, undif- 

7: 198 

ferentiated, bitegmic, only in ripe seed the 
inner integument wholly or largely sup- 


Add under Viola the following species: 
Viola rheophila Okamoto, Bull. Osaka 
Mus. Nat. Hist. no. 37 (1984) 4-15, 3 fig. 
Distr. Malesia: Borneo (Kalimantan 


compiled by 


Families and higher taxa have been entered under their name. 

Names of families which have been revised in volumes 4-10 have been entered and are printed in bold 
type, so that as far as this is concerned this index is complete for all preceding volumes as well. 

Suprageneric epithets have been entered under the family name to which they belong preceded by the 
indication of their rank (subfamilies, tribes, etc.). 

Infrageneric epithets have been entered immediately under the generic name to which they belong, pre- 
ceeded by the indication of their rank (subgenera, sections, series, etc.). 

Infraspecific epithets have been entered under the specific name to which they belong preceded by the 
indication of their rank (subspecies, variety, forma, etc.). 

Epithets of new names and new combinations have been printed in bold type, synonyms in italics. 

Page numbers in bold type denote main treatment; an asterisk behind a page number denotes the 
presence of a figure of the concerned texon; 'map' printed behind a page number denotes that a map of the 
concerned taxon is present on that page. 

Some minor printing errors in plant names have been corrected. 

Of synonyms with a double authority, the latter has not always been cited in full. The full authority can 
easily be derived from the text. 

Abies dammara Desf. 433 
Aceraceae 4: 3-4, 592; 6: 

915; 7: 820; 8: 549 
Acioa Aubl. 639 

heieropeiala (ScorL ex 
King) Kosierm. 677 
malayana Kosterm. 677 
pcrcoriacea Kosterm. 678 
Acioa aucl. 675 
Acmopyle 343, 352, 354 
Actinidiaceae s.str.4: 37-39 
Adenia 166 
Afrocarpus 354 
Afrolicania Mildbr. 645 
Agathis Salisb. 339,341-343, 
347, 354,420,421, 429, 
430', 431 map 
sect. Agathis 431 
sect Macrobracteaiae Mcijer 

Drees 431 
sect. Microbracleatae Mcijer 

sect. Prismatobractcatae 

Meijer Drees 431, 440 
sect. Rostrata dc Laub. 431, 

alba Foxw. 433 
fl/6flauct. 435,437 
beccarii Warb. 433 
beckingii Mcijer 435 
bornccnsis Warb. 340, 34 1 , 
430*. 432*. 433, 434* 
cclcbica (Koord.) Warb. 433. 
434. 435 map 
ssp. ceiebica Vcldk. & 
Whitm. 435 

(Agathis ceiebica) 

ssp. flavescens (Ridl.) 
Veldk. & Whitm. ex 
Veldk. & de Laub. 438 
dammara (Lamb.) Richard 
433, 434 

ssp. dammara aucl. 435, 

ssp. dammara VJhilm. 433 

ssp. flavescens (Ridl.) 
Whitm. 438 
dammara auct. 435, 437 
endcrtii Mcijer Drees 439, 

440* map 
flavescens Ridl. 340, 341, 

433, 438 
hamii Meijer Drccs 435 
kinabalucnsis de Laub. 433, 

438 map, 439 
labiilardieri Warb. 436 map, 

440, 441* 
latifotia Mcijer Drees 433 
Icniicula de Laub. 433, 

436*, 437 map 
loranthifolia Salisb. 433 
loranthifolia auct. 435 
macrostachys Warb. 433 
motleyi (Pari.) Diimmcr 

orbicuia de Laub. 433, 437* 

palmerstonii (F.v.M.) Bailey 

philippincn.sis Warb. 433, 

437. 438 map 
regia Warb. 437 
rhomboidales Warb. 433 


robusia (Moore) Bailey 442 
ssp. nesophila Whitm. 

var. nesophila auct. 442 
var. robusia Whitm. 442 
spathulata dc Laub. 433, 
435, 436 map 
Agonandra 31-34 

raccmosa 33 
Ailanthus intcgrifolia Larnk 718 
Aizoaceae 4: 267-275; 6: 

Alberlisia Becc. 160-162, 164, 
167-169, 171, 179 
crassa Forman 179, 181 
mccistophylla Forman 181 
mcgacarpa Diels ex Forman 

159, 179, 181 
papuana Becc. 166, 179, 
180*, 181 
Alc.imandra Dandy 565, 566. 
567, 569 
cathcariii (Hk.f. & Th.) 
Diindy 569 
Alismataceae 5: 317-334; 

6: 915, 9: 553 
Alismauilcs 109 
Allantospcrmum Forman 621, 

Alnus 338 
Alpam Rhccde 65 
Alscuosmia 335 
AlMCUosmiaccac 335-336 
Aisoinitra macrcxarj^a (Bi.) 
Rocm. 29 



Flora Malesiana 

[ser. I, vol. 104 

Amaranthaceae 4: 69-98, 

593; 5: 554; 6: 915; 

7: 820; 8: 549 

Ambora ficus Tul. 326 
Amborclla 261 
Amorpha pedalis Blanco 488 
Anacardiaceae 8: 395-548; 

9: 553; 10: 679-681 
Anaoirdium occidcntale 352 
Anacolosa Bl. 1,3-6, 23 
arbor ea K. & V. 25 
cauliflora Sleum. 25 
celebica Valet. 25 
frutcscens (Bl.) Bl. 2, 24*, 

25, 26* map 
heptandra Maing. ex Mast. 25 
luzoniensis Mcrr. 25 
maingayi Mast. 22 
papuana Schelle.nb. 25, 26 
Zollinger I Baill. 25 
Anamirta Colcbr. 157, 158, 

160-162, 166-168, 170, 
171, 211 
cocculus (L.) Wight & Am. 
159, 165. 166, 212*, 
213, 214* 
flavescens (Lamk) Miq. 210 
jucunda Micrs 213 
lemniscata Miers 210 
loureiriPicTTC 210 
luctuosa Miers 210 
Anaxagorea A. St. Hil. 605 
Ancistrocladaceae 4: 8-10; 

5: 553 
Ancistrocladus pentagynus 

Warb. 612 
Androglossum reliculaium 

Champ, ex Benth. 685 
AndrurisSc\\\\s\09, 110 

andajensis (Becc.) Schltr 118 
anisophylla Cicscn 112, 118 
australasica (Hemsl.) Giesen 

bur uensis J. J. Smith 118 
celebica Schltr 118 
clemensae (Hemsl.) Giesen 

112, 118 
clemensae (Hemsl.) Giesen 

var. borneensis Giesen 118 
crinila (Becc.) Schltr 1 17 
elegans G'lescn 118 
gracillima Giesen 1 17 
javanica Giesen 118 
loheri Giesen 117 
nana (Bl.) Giesen 117 
palawensis Tuyama 118 
/ene//a Schltr 118 


viiiensis (A.C.Smith) Giesen 

wariana Schltr 1 17 
Angelesia Korth. 645 
papuana Baker f. 65 1 
splendens Korth. 646 
Anisadcnia 609 
Anisophyllaea gaudichaudiana 

Baill. 253 
Annonaceae 63, 64, 261, 605, 

Anthobembix Perkins 306, 308 
brassii A.C. Smith 315 
dentatus \ a\Q.iox\ 310 
hospilans (Becc.) Perkins 311 
ledermannii Perkins 319 
moszkowskii Perkins 3 1 1 
myrtifolia A.C. Smith 313 
oligantha Perkins 315 
parvifolia Perkins 315 
Antidesma ghacsembilla Gaertn. 
pcntandrum (Blco) Merr. 52 
Ami taxis Miers 172 
calocarpa Kurz 174 
cauliflora (Miers) Dicls 174 
fasciculata Miers 174 
longifolius (Decne ex Miq.) 
Miers 174 
Apama Lamk 62, 63, 65 
affinis Weisse 79 
brevipes Weisse 79 
corymbosa (Griff.) Willd. ex 

Soler. 78 
macranlha Weisse 81 
tomeniosa Engl, ex Soler. 
var. lanuginosa (Hk.f.) 
K. & G. 79 
iAp/zy/Ze/a Champion 110 

erubescensCUdsn^). 114 
Aponogetonaceae 4: 11—12; 

5: 553; 7: 213 
Aptandra 3, 4 
Aptandropsis 4 
Aquifoliaceae 145 
Aquilegia 165 
Ardbidopsis 543 
Arabis 54 1 
Araliaceae 9: 1-105, 553; 

10: 716 
Araucaria Juss. 337, 339, 421, 
423 map 
sect. Araucaria 422, 423 
sect. Bunya Wilde & Eames 

sect. Columbea All 
sect. Colymbea Endl. 422 


sect. Eutacta Endl. 422, 423, 

sect. Euiassa (Salisb.) Benth. 

& Hook. 425 
sect. Intermedia White 423 
subg. Colymbea Antoine 423 
subg. Eutacta (Link) Antoine 

beccarii Warb. 427 
bidwiiiii 420, 423 
cunninghamii Ait. ex D. Don 
340, 347, 425 

var. papuana Laut. 426*, 
427 map, 428* 
cunninghamii auct. non Ait. 

hunsteinii K.Sch. 339, 347, 

420, 423, 424 map 
klinkii Laut. 423, 424 
schumanniana Warb. 424 
Araucariaceae 338, 339, 343, 

347, 354, 419-442 
Araucariacites 338 
Arcangelisia Becc. 158, 160, 

161, 166-171, 209 
flava(L.) Mcrr. 160, 210, 

inclyta Becc. 210 
lemniscata (Miers) Becc. 210 
loureiri (Pierre) Dicls 210 
tympanopoda (Laut. & 

K. Sch.) Diels 210, 211 
Archichlamydeae 127 
Ardisia glabra (Thunb.) DC. 134 
Aristolochia L. 53-55, 57-59, 

61-64,83, 166 
acuminata Lamk 94 
barbata 55 
brasiliensis Mart. & Zucc. 

clematitis 59 

coadunata Backer 57-59, 84, 
86, 95, 96* 

var. bosschai Backer 95 

var. coadunata 95 
crassinervia Schmidt 58, 82*, 

85, 87, 104* 
curtisii King 84, 86, 89 
decandra Ding Hou 57-59, 

64, 83, 85, 87, 100, 

101*, 102* 
deltantha F.v.M. 97 
dictyophlebia Merr. & Perry 

dielsiana Schmidt 57, 61, 64, 

84, 85, 88, 105 
elegans Mart. & Zucc. 55, 



Index to scientific plant names 



engleriana Schmidt 85, 88, 

foveolata Merr. 55, 58, 85- 

87, 91 map 
gaudichaudii Duchartre 56*, 

57, 84, 88, 107 
gigantea Mart. & Zucc. 64 
glaucifolia Ridl. 55, 85, 86, 

gracilifoUa Schmidt 97 
grandiflora Sw. 57, 59, 62, 

griffiihii Duchartre 59, 91 
hastata Jack 88 
hastata (non Jack) Klotzsch 

humilis Merr. 55, 85, 87, 

imbricaia Mast. 103 
indica L. 99 
indica (non L.) BIco 94 
indica (non L.) L. 97 
jackii Steud. 55, 58, 59, 84, 

86, 87, 88, 89* 
japonica Miq. 94 
kaempferi WiUd. 92 
kankauensis Sasaki 92 
kaoi Liu & Lai 91 
klossh Ridl. 85, 86, 93 
kwangsiensis Liang 91 
lautcrbachiana Schmidt 108 
ledermannii Schmidt 106, 

107 map 
Icuconeura 62 
leytensis Merr. 84, 85, 87, 

linnemannii Warb. 55, 85, 

88, 107 
longifolia Roxb. 94 
macgregorii Merr. 55, 56*, 

57, 85, 87, 103 
maurorum 58 
megalophylla K.Sch. 94 
memhranacea Merr. 103 
mindanaensis Warb. 94 
minuiiflora Ridl. ex Gamble 
55. 85-87, 93 

var. dolahrata Gamble 93 
moluccana Duchartre 94 
momandul K. Sch. 58. 59, 

82*, 85, 86, 88, 97 
naviculilimba Ding Hou 85, 

87. 100* 
novogumcCnsis Schmidt 108 
ornilhrjcephala 55 
papillifolia Dmg Hou 58, 84. 

87. 98*. 99 
pcliata 59 


philippinensis Warb. 60*, 

66, 84, 87, 102 
piihecurus Ridl. 97 
ramosii Merr. 92 
ringens Vahl 55, 64 
roxburghiana Klotzsch 94 

6 angustifolia Duchartre 

ssp. kankauensis (Sasaki) 
Kitamura 92 
roxburghiana (non Klotzsch) 

Warb. 107 
rumphii KosteleL?Jcy 85, 86, 

97, 101* 
samarensis Merr. 85, 87, 

schlechteri Laut. 58, 59, 85, 

88, 97, 105, 106, 107 
sericeaBlco55, 57, 58, 61, 

66, 85, 87, 103 
singalangensis Korth. ex 
Ding. Hou 61, 84, 86, 

89, 90* 

tagala Chamisso 54, 55, 56*, 

57, 63, 85-88, 94 
tagala (non Chamisso) Halus. 
var. hankaoensis 92 
var. kankauensis (Sasaki) 
Yamazaki 92 
timorensis Dccne 94 
Lranstillifera Ding Hou 60*, 

84, 87, 99 
iripariiia Backer 88 
ungulifolia Mast. 88 
zollingeriana Miq. 57, 85, 
86, 92 
Aristolochiaceae 53-108, 

126, 166 
Aristolochiales 567 
Aromadendron Bl. 567, 568, 
ashionii Dandy ex Cockbum 

borneensis Dandy ex Cock- 
bum 579 
elegans Bi. 577 

var. glauc.a (Korth.) 
Dandy 577 
glaucum Korth. 577 
nutans Dandy 577 
oreadum (Diels) Kanch. & 
Hatus. 583 
Arorrtadendrum 568 
Aromoracia rusticana G.M. & 

Schcrb. 543 
Asarum 57 
virginitum 62 

Ascarina J.R. & G. Forst. 123- 
129, 137, 140 map 
sect- Ascarina 137, 139, 140 

sect. Madascarina Leroy & 
Jeremie 123, 137, 140 
diffusa A.C. Smith 139, 

lanceolata (non Hk.f.) auct. 

lucida 124, 139 
maheshwarii Swamy 139, 

140, 141 
philippinensis C.B. Rob. 

138*, 139, 141 
reticulata Merr. 139 
rubricaulis 127 
subsessilis Verdcourt 139, 
Ascarinopsis 123, 128 
Ascarinopsis Humb. & Capuron 

123, 128, 137, 140 map 
As hto n s Aromadendron Mcijcr 

Asiphonia Griff. 65 
piperiformis Griff. 78 
sp. Griff. 78 
Aspidocarya 162 

dissitiflora Laut. & K. Sch. 

hirsuta^cjcc. 193 
kelidophylla Laut. & 

K. Sch. 253 
uvifcra Hk. f. & Th. 189* 
Aihcrospcrmataceae 261 
Athcrospermoxylon 256 
Athrotaxis 339 
Au-iaecarpum 158 
Atroxima 456 

Atuna Rafm. 636-642, 665 
cordata Cockburn ex Prance 

636, 666 map, 667 
data (King) Kostcrm. 669 
elliplica Ko.stcrm. 666 
excelsa (Jack) Kostcrm. 636, 

indica (Bald.) Kostcrm. 666 
latifolia (Hcnd.) Kostcrm. 

lalifrons (Kostcmi.) Prance & 

While 666 map 
nannodes (Kostcrm.) 

Koslerm. 666. 667 map 
pcnangiana (Kostcnn.) 

Ko.stcrm. 666. 667 map Rafin. 666. 669 
ssp. (Jack) I'rancc 
668*. 669. 670 map 


Flora Malesiana 

[ser. I,vol. 104 

(Atuna racemosa) 

ssp. racemosa 669, 670 
scabra (Hassk.) Koslerm. 

travancorica (Bedd.) Kosterm. 

villamilii (Merr.) Kosterm. 
Aiunus Rumph. 665 
alba Rumph. 669 
Austrocedrus Florin & Boutelje 

Austrotaxus 339, 348 

Badiera (non DC.) Hassk. 459, 
pulchra (Hassk.) Hassk. 468 
venenosa (Poir.) Hassk. 468 
Bafodeya Prance 642 
Balanophoraceae 7: 783—805; 

8: 549; 9: 554 
Balantium Desv. ex Buch.-Ham. 

Balsaminaceae 539 
Bania Becc. 183 

thyrsiflora Becc. 183 
Banisterodes O.K. 493 
affine (Miq.) O.K. 503 
ellipilcum (Korth. ex Miq.) 

O.K. 530 
excelsum (Bl.) O.K. 501 
glaucum (Wall, ex Hassk.) 

O.K. 527 
griffiihii (Hk.f. ex Benn.) 

O.K. 513 
insigne (Benn.) O.K. 537 
/on^//b//a(Bl.)O.K. 514 
maingayi (Benn.) O.K. 524 
obscurum (Benn.) O.K. 537 
rufum (Benn.) O.K. 505, 

stipitatum (Benn.) O.K. 535 
vitellinum (Bl) O.K. 515 
Bannisierioides 493 
Barbarea vulgaris R. Br. 545 
Bamharlia Gleason 493 
Basellaceae 5: 300-304 
Baiania Ualus. 172 

insignis Hatus. 174 
Baterium Miers 183, 184 

validum Miers 1 84 
Bat(id)aceae 5: 414-415; 6: 

Benneiiiodendron leprosipes 

(Clos) Merr. 715 
Bcrberidaceae 163 
Berbcridales 163 

Betulaceae 5: 207-208; 6: 

Bignoniaceae 8: 114-186; 

9: 554; 10: 716 
Bixaceae s.str. 4: 239-241 
Bladhia glabra Thunb. 1 34 
Blumia Nees 569 

candollii (Bl.) Necs 582 
Blyxa alternifolia (Miq.) Hanog 
aubenii Rich. 717 

var. echinosperma (Clarke) 
Cook &Luond 717 
echinosperma (Clarke) Hk.f. 

japonica (Miq.) Maxim, ex 
Aschers. & Gurke 717 
var. alternifolia (Miq.) 
var. japonica 717 
leiosperma Koidz. 717 
Boraginaceae 29 
Borneo Aromadendron Meijer 

Bracea King 29 
Brachynema 3, 4 
Braganlia Lour. 65 

affinis Planch, ex Rolfe 79 
blumii Lindl. 79 
brevipes Merr. 79 
corymbosa (non Griff.) 

F.-Vill. 103 
corymbosa Griff. 78 
macranlha Boerl. 81 
melastomaefolia Duchartre 

tomentosa Bl. 79 

var. lanuginosa Hk. f. 79 
Brassica L. 541, 543, 545 
besseriana Andrz. ex Trautv. 

campestris L. 543 
chinensis L. 543 
integrifolia (West) Rupr. 546 
juncea (L.) Czem. 544*, 545 
napus L. 543 
oleracea L. 543 
rapa L. 543 
rugosa Prain 543 
Bredemeyera 456, 457, 465 
sect. Melchiora Steen. 465 
floribunda 457 
papuana Steen. 465 
Brewsiera M.J. Roemer 622, 
crenaia M.J. Roemer 625 
Brongniartia Bl. 287 

coriacea Bl. 298 
Brownetera L.C. Rich. 356 

Bruguiera gymnorrhiza (L.) 

Savigny 717 
Bubbia 123 
Burmanniaceae 4: 13-26, 

592; 5: 553; 7: 820; 

9: 554 
Burseraceae 5: 209-296, 

567; 6: 917; 7: 820; 

9: 555; 10: 715 
Butomaceae 5: 118-120, 

Byblidaceae 7: 135-137 

Callitrichaceae 4: 251-252 
Callitris sp. 453 
Callitroideae 444 
Calocedrus 444 
Camelina 541 
Campanulaceae 6: 107—141, 

928; 8: 549; 9: 556 
Canariopsis anguslifolia Bl. ex 

Miq. 715 
Canarium angustifolium Miq. 

Candjera 5 1 

Cannabinaceae 4: 222-223 
Cansiera 50, 52 

zyziphifolia 50 
Cansjera Juss. 31-35, 48, 49 
grossularioides Blco 52 
lanceolala Bth. 50 
leplostachya Bth. 31, 33, 34, 

48*, 49 map, 51* 
leplostachya Koord. 35 
malabarica Lamk 50 

var. 6 50 
manillana Bl. 36 
monostachya (Willd.) 

M. Roemer 50 
parvifolia Kurz 48*, 49 map 
penlandra Blco 52 
polystachya (Willd.) 

M. Roemer 50 
rheedii J.F. Gmelin 31-33, 

48*, 49 map, 50* 
rheedii Blco 52 
scandens Roxb. 9, 50 
timorensis Decne 5 1 
zizyphifolia Griff. 50, 51 
Cappar(id)aceae 6: 61-105; 

7: 822 
Caprifoliaceae 4: 175-194; 
6: 928; 9: 556; 10: 335 
Capsella Medicus 541, 543, 
545, 549 
bursa-pastoris (L.) Medicus 


Index to scientific plant names 


Cardamine L. 541, 543, 545, 
africana L. 541, 551 
ssp. borbonica (Pers.) 
var. papuana Laul. 55 1 
alligena O.E. Schulz 551, 

borbonica Pers. 551 
decurrens (Bl.) Z. & M. 554 
flexuosa With. 551, 554 
flexuosa auct. 554 
hirsuta L. 551, 554 
javanica (Bl.) Miq. 552 
keysseri O.E. Schulz 551, 

papuana (Laut.) O.E. Schulz 

551, 552* 
re ge liana Miq. 554 
sublyrata Miq. 559 
Cardiopteridaceae 7: 93—96; 

10: 166, 253, 716 
Cardioptcris Royle 166, 716 

moluccana Bl. 19, 253 
Carex bilateralis Hayata 716 
Caricaceae 61 
Carpolobia 456, 457 
Carronia F.v.M. 158, 160, 162, 
167, 168, 170, 172, 183 
thyrsiflora (Becc.) Dicls 
182*, 183 
Carson's Magnolia Meijer 571, 

Cascaria flavovirens Bl. 716 

pallida Craib 716 
Casuarinaccae 453 
Cathedra 3, 4 
Cayloniales 338 
Caytonipollcniies 338 
Celastraceae 6: 227-291, 
389-421,930; 10: 
539, 716 
Cclaslrales 145, 629 
Celasiranac 151 
Ccliis paniculaia (Endl.) Planch. 

Ccntrolepidaceae 5: 421-427 
Ccphaiotaxaccac 343, 347 
Ccphaloiaxus 354 
celebica Warb. 349 
mannii (non Hk.f.) F*rilzcl ex 

Dicls 349 
sumatrana Miq. 349 
Ceramium Bl. 65 

lomenlosum fil. 79 
Ceratophyilaceae 4: 41-42 
CV.ri<jps dctandra 7 1 8 
ChamaehuxiLS (DC.) Spach 459, 


arillaia (D. Don) Hassk. 469 
pulchra (Hassk.) Hassk. 468 
venenosa (Poir.) Hassk. 468 
sub var. elliptica Miq. 468 
subvar. obovata Miq. 468 
var. apiera Miq. 468 
var. gracilis Miq. 468 
var. minor Miq. 468 
var. robusla Miq. 468 
Champaca Adans. 598 
lurbinata Nor. 583 
Champereia Griff. 31-34, 35 
cumingiana (Baill.) Merr. 36 
gaudichaudiana (Baill.) Tiegh. 

griffithii Planch, ex Kurz 36 
lanceolata Merr. 36 
manillana (Bl.) Merr. 31, 34, 
35, 36*, 37*, 38 map, 40* 
oblongifolia Merr. 36 
perrolteiiana Baill. 52 
platyphylla Merr. 36 
Champereya 35 

gnetocarpa Kurz 36 
griffiihiana Planch, ex Kurz 
Chandlera 158 
Chasmanihera 164 
Chaunochiton 3, 4 
Cheirolepidaceae 338 
Chenopodiaceae 4: 99—106, 
594; 6: 932; 8: 549; 9: 
Chlaenandra Miq. 158, 161, 

162, 167, 168, 170, 171, 
ovaia Miq. 159. 186*, 187, 
Chloranthaceae 123-144 
Chloranihales 127 
Chloranlhus Swariz 123-128, 
129. 139. 143, 144 
brachysiachys Bl. 

var. melanocarpus Ridl. 
brachysiachys scnsu Benlh. 

chincnsis 124 
denliculalus Cordemoy 1 34 
elalior R.Br, ex Sims 131 
creclus (Buch.-Ham.) Verde. 
124. 127. 128. 130*. 
131. 132*. 133*. 136 
ereclusSwccl 130 
fortunci 127 
glabcr scnsu Backer & 

Bakh.f. 136 
glaber (Thunb.) Makino \?A 


hainanensis Pei 1 36 
henryi Hemsley 130, 133 
inconspicuus Svj2iT\z 133 
inconspicuus (non Swartz) 

Blco 131 
indicus Wight 133 
japonicus 127, 130 
monander R. Br. ex Sims 

multisiachys Pei 128 
obtusif alius Miq. 133 
officinalis Bl. 124, 127, 

128, 131, 132 
oldhamii Solms-Laub. 133 
oldhamii (non Solms-Laub.) 

Merr. & Quis. 133 
philippinensis Merr. 133 
salicifolius Presl 131 
serratus 127 
spicalus (Thunb.) Makino 

127, 128, 130, 132 
veriicillatusMen. 133 
Chrysobalanaceae 635-678 
tribe Chrysobalaneae 635 
tribe Couepieae 636 
tribe Hinelleae 636 
tribe Parinarieae 635 
Chrysobalanus L. 635-637, 

639, 640, 642, 643 
ciliatus Korth. ex Miq. 674 
icaco L. 643, 644* map 
splendens Korth. ex Miq. 

Cissampelos L. 157, 158. 160- 

162. 165. 167. 169-171, 

convolvulacea Willd. 236 

var. hirsuta (DC.) Hassk. 
cumingiana Turcz. 236 
discolor DC. 236 

var. cardiophylla A. Gray 
hernandifolia 245 
hernandiifotia Willd. 245 
insularis Makino 240 
ovala Poir. 217 
pareiraL. 165 

var. hirsuia (Buch. ex 
DC.) Forman 235*. 

var. orbiculaia (DC.) Miq. 

var. peltata Schcff. 236 

var. typica Diels 236 
parcira (non L.) Ridl. 229 
psilophylla Prcsl 25 1 


Flora Malesiana 

[ser. I, vol. 104 

Citronella suaveolens (Bl.) 

Howard 717 
Cladopus nymani H. M611. 717 
Classopollis 338 
Clethraceae 7: 139-150 
Clypea BI. 243 

acuminalissima Bl. 252 
capitata Bl. 252 
corymbosa Bl. 249 
discolor BI. 245 
glaucescens Decne 245 
tomentosa Bl. 229 
venosa Bl. 248 
Coccomelia Ridl. 645 
CocculusDC. 157, 158, 161, 
162, 165, 169, 231 
angustifolius Hassk. 234 
bantamensis Bl. 199 
blumei Boerl. 223 
brachysiachyus DC. 217 
celebicus Boer). 175 
cinereus Zoll. & Mor. 229 
coriaceus Bl. 199 
corymbosus Bl. 229 
crispus [non (L.) DC] Hassk. 

cuspidatus V^ all. 2\9 
cynanchoides PresI 233 
elegans (Ridl.) Ridl. 233 
ferrandianus Gaudich. 233 
flavescens (Lamk) DC. 210 
flavicans Wall. 253 
forsteri DC. 245 
glaucescens BI. 230 
^/aucwi (Lamk) DC. 229 
incanus Colebr. 229 
japonicus (Thunb.) DC. 245 
var. limoriensis DC. 245 
kunslleri King 231 
lanuginosus Bl. 229 
laurifolius DC. 160, 166, 
169, 171,231,232*, 
233, 234 
var. angustifolius (Hassk.) 

Boerl. 233 
var. triplinervis Boerl. 233 
leptostachyus DC. 217 
limacia DC. 223 
longifolius Decne ex Miq. 

lucidus Teijsm. & Binn. 174 
macrocarpus W. & A. 230 
mollis Hk.f. & Th. 233 
orbiculalus (L.) DC. 159, 
160, 162, 170, 172, 
231, 232* 
ovalifolius (Vahl ex Pers.) 

DC. 233 
populifolius DC. 213 


rimosus Bl. 207 
sarmenlosus Dicls 233 
var. stenophyllus Merr. 
tr iandr us CoXchx. 185 
/r//?o/'U5DC. 219, 233 
/n7(3feu5 (Thunb.) DC. 159, 
163, 165,231,233,234 
velutinus Wall. 223 
Cochlearia officinalis L. 543 
Cochlospermaceae 4: 61—63 
Coffea anguslifolia Roxb. 717 
Columbea All, Al^ 
Colymbea Spreng. 422 
Combretaceae 4: 533-589; 
5: 564; 6: 932; 7: 823 
Comesperma 456, 461, 465 
sect. Prosthemosperma 
F.v.M. 465 
Coniferales 337-453 
Connaraceae 5: 495-541; 
6: 933; 7: 823; 8: 549; 
9: 557 
Convolvulaceae 4: 388- 
512,599; 5: 558; 6: 
936; 7: 823; 9: 558; 
10: 29,716 
Cordai tales 126 
Cornaceae 8: 85-97 
Coronopus 541 
Coryneliales 353 
Corynocarpaceae 4: 262— 
264; 5: 557; 6: 941 
Corytholobium Mart, ex Benth. 

Coscinium Colebr. 160-162, 
166-168, 170, 171, 215 
blumeanum Micrs ex Hk.f. & 
Th. 212*, 215, 216, 
blumeanum (non Miers ex 
Hk.f. &Th.) Miq. 216 
var. epel latum Boerl. 216 
fenestratum (Gaertn.) Colebr. 
160, 165, 212*, 215, 
var. macrophyllum 
var. ovalifolium 
maingayi Pierre 216 
mangayi 216 
miosepalum Diels 216 
peltatum Merr. 216 
wallichianum Miers 215 
Couepia 640 
Coula 3, 4 

Crassulaceae 4: 197-202; 

9: 558 
Cri.spiloba Stccn. 335 
Croialaria duboisii Lev. 469 
Cruciferae 541-560 
tribe Arabideae 543 
tribe Brassiceae 543 
tribe Heliophilcae 543 
tribe Lepidicae 543 
tribe Stcnopctaleae 543 
Cryphaea Buch.-Ham. 129 

erecta Buch.-Ham. 131, 132 
Crypteroniaceae 8: 187-204 
Cryplocarya reticulata BI. 715 
Ctenolophon Oliver 29, 629 
englerianus Mildbr. 630, 631 
grandifolius Oliver 631 
parvifolius Oliver 630, 631, 

632*, 633* map 
philippinensis Hall.f. 630, 
Ctenolophonaceae 607, 609, 

Cucurbitaceae 29, 166, 253 
Cupressaceae 338, 339, 343, 

Curupira 3, 4 

Cyclandrophora Hassk. 642, 665 
asperula (Miq.) Prance ex 

Kosterm. 671 
elata (King) Kosterm. 669 
excelsa (Jack) Kosterm. 671 
glaberrima Hassk. 669 
lalifolia (Hend.) Prance 666 
laurina (Gray) Kosterm. 669 
nannodes (Kosterm.) 

Kosterm. & Prance 667 
penangiana Kosterm. & 

Prance 667 
scabra (Hassk.) Kosterm. 669 
villamilii (Merr.) Prance ex 
Kosterm. 671 
Cyclea Am. ex Wight 160-162, 
165, 167, 169, 170, 172, 
acuminatissima Merr. 239 
apoensis Yamamoto 24 1 
atjehensis Forman 237, 238*, 

barbata Miers 159, 136, 139, 

242, 243, 253 
barbata Craib 239 
caudata Merr. 239 
cauliilora Merr. 237, 241 
ciliata Craib 242 
elegans King 187, 237, 239 
insularis (Makino) Hatus. 
237, 240 
var. luxurians Hatus. 240 


Index to scientific plant names 



kinabaluensis Forman 159, 
237, 239 
var. hispida Forman 239 
korihalsii Diels ex Norman 

laxiilora Miers 237, 239, 

241, 242, 243 
merrillii Diels 237, 240 
peliata [non (Lamk) Hk. f. & 
Th.] Becc. 241 
var. arnouii Miers 241 
peliata [non (Lamk) Hk. f. & 

Th.] Miq. 163, 242 
percgrina Miers 237, 239, 

241. 242 
robusia Becc. 237, 240 
scyphigera Suesseng. & 
Heine 239 
forma anguslifolia 

Suesseng. & Heine 239 
tomcntosa 243 
tonkinensis (non Gagnep.) 

Yamamoto 239 
wallichii Diels 242, 243 
Cyclodiscus tomentosus 

Klolzsch 79 
Cyperaceae 7: 435—753, 

823; 9: 107-187,560; 
10: 716 
C>perus diaphanus Schrader ex 
R. & S. 
var. latespicalus (Boeck.) 
Kern 716 
CyrillopsisKuhlm. 621,622 

Dacrycarpus (Endl.) dc Laub. 
337, 338,343, 347, 351, 
353-355. 374, 376 map 
cincius (Piiger) de Laub. 376, 

383*, 384 map 
compaclus (Was.scher) de 
Laub. 376, 382* map, 
383*. 384 
cumingii (Pari.) de Laub. 

376, 381 map 
dacrydiifoUa (Was-schcr) 

Gausscn 383 
cxpansus dc Laub. 342. 376, 

381, 382 map 
imbncatas (Bl.) dc Laub. 
340, 341. 376 
var. turvulus (Miq.) dc 
Laub. 340, 341. 377. 
379 map, 380* 
var. imbncatus 377 map 
var. palulas dc l^ub. 377 
map. 378*. 379 

(Dacrycarpus imbncatus) 

var. robustus de Laub. 
377, 379 map 
kawaii (Hayata) Gaussen 

kinabaluensis (Wasscher) de 

Laub. 376, 381 map 
leptophylla (Wasscher) 

Gaussen 371 
steupii (Wasscher) de Laub. 

340, 376, 380 
steupii (non de Laub.) de 

Laub. 379 
Dacrydium Soland. ex Forsl. f. 
337, 338, 342, 343, 347, 
352-355, 360, 362 map 

balansae 365 

beccarii Pari. 362, 366, 367 
var. rudens de Laub. 368 
var. subelatum Comer 363 

comosum Comer 363, 370, 
371 map 

cornwalliana de Laub. 340, 

362, 364 map, 366* 
cupressinum 365 

datum (Roxb.) Wall, ex Hk. 

341, 362, 363. 364 map 
ericoides de Laub. 340, 

341, 363, 371 map 
falciforme (Pari.) Pilgcr 372 
falciforme [non (Pari.) Pilgcr] 

Foxw. ex Mcrr. 373 
falciforme [non (Pari.) Pilgcr] 

Laut. 373 
gibbsiae Siapf 363. 369 
gracilis de Laub. 363, 367 

junghuhnii Miq. 363 
leptophyllum (Wa.sschcr) 

dcLaub. 363,371 
magnum de Laub. 363, 368 

medium dc Laub. 339-341, 

363, 368 map 
micropedunculatum 340 
nidulum de Laub. 362. 365 

var. araucarioides dc Laub. 
novo-guincensc Gibbs 362, 

364 map 
pctlmatum de Laub. 340. 
.34 1 . 362, 364 map 
var. robusium dc Laub. 
pterrii Hickcl 363 
spathoidcs dc I-aub. 340, 363, 
367 map. 368 


xanihandrum Piiger 363, 
369*, 370* map 
Dactyladenia 639 
Dahuronia Scop. 645 
Dammara Link 429 

alba Rumph. ex Hassk. 435 
var. alba Hassk. 435 
var. celebica Hassk. 435 
celebica Koord. 435 
loranthifolia Link 433 
motleyi Pari. 393 
orienialis Lamb. 433 

var. alba Knight ex Henkel 

& Hochst. 435 
var. orienialis Carr. 433 
var. pallens Carr. 435 
palmerstonii F.v.M. 442 
robusia Moore ex F.v.M. 

rumpfii auct. 437 
rumphii Presl 435 
Daphnandra Perkins 256, 261, 
263, 265 
aromadca 261 

novoguineensis Perkins 266 

perkinsiae Gilg & Diels 266 

Daphne monosiachya Willd. 50 

polysiachya Willd. 50 
Datiscaceae 4: 382-387; 7: 

Davisicarpum 158 
Decussocarpus de Laub. 389 
sect. Dammaroides (Bennett) 

de Laub. 390 
.sect. Decussocarpus de Laub. 

maximus de Laub. 394 
motleyi (Pari.) de Laub. 393 
viiiensis (Seem.) dc Laub. 

wallichianus (Presl) dc Laub. 
Dcgcncriaceac 562 
Dichapetalaceae 5: 305-316. 

567; 6: 941; 7: 823 
Diemenia Korth. 648 

racerrwsa (Korlh.) Miq. 649 
Dillcnia 123 
Dilleniaceae 4: 141-174; 

5: 557; 7: 824 
Dimocarpus longan 159 
Diogoa 3, 4 
Dioscorca 166 

aculcata (non L.) Zoll. 199 
spiculala Bl. 199 
slcnomcridora Pram ik Hurk. 


Flora Malesiana 

[ser. I, vol. 104 

Dioscoreaceae 4: 293-335; 

5: 557; 10: 253 
Dioscorcophyllum cumminsii 

Diploclisia Miers 158, 160, 
169, 170, 172, 229 
glaucescens (Bl.) Diels 230 
glaucescens (non (King) 

Diels) sensu Forman 23 1 
kunstleri (King) Diels 228*, 

macrocarpa (W. & A.) Miers 
Dipsacaceae 4: 290-292; 

5: 557 
Dipterocarpaceae 9: 237—552; 

561; 10: 665, 716 
Dipierocarpus comutus Dyer 665 
Discogyne Schitr 622 
papuana Schitr 627 
Dolichandrone spalhacea (L.f.) 

K.Sch. 716 
Dombeya Lamk. 422 
Dor>'phora 261 

aromatica 261 
Drebbelia Zoll. 6 

subarborescens Zoll. 7 
Droseraceae 4: 377—381; 

5: 557; 6: 943; 9: 562 
Dryadodaphne S. Moore 255- 
257, 262, 263 
celastroides S. Moore 266 
crassa Schodde 264*, 265, 

novoguineensis (Perkins) 
A.C. Smith 261, 265, 
266, 267 
ssp. novoguineensis 266, 
var. macra Schodde 

var. novoguineensis 
266, 267 
ssp. occidentalis 
Schodde 265, 267 
novoguineensis (Perkins) 
A.C. Smith p.p. 265 
Drymis-leaved Magnolia Meijer 

571, 573 
Dugoriia Scop. 654 
Dulacia 4 

Durandea Planch. 608, 609 
jenkinsii (F.v.M.) Siapf 612 
magnifolia Stapf 615 
pallida K.Sch.eW 
pentagyna (Warb.) K. Sch. 
var. rolundaia (K. Sch.) 
Laul. 612 


robinsonii (Merr.) Hall.f. 612 
rolundaia K. Sch. 612 

Elaeagnaceae 151-156 

Elaeagnales 1 5 1 
Elaeagnus Toum. ex L. 151, 
seel. Elaeagnus 152 
sect. Sempervirentes Serv. 

anguslifolia (non L.) Blco 

arborea Roxb. 
var. dendroidea 
Schlechtend. 153. 
conferta Roxb. 152, 153, 
ssp. dendroidea 

(Schlechtend.) Serv. 
ssp. euconferia Serv. 153 
ssp. javanica (Bl.) Serv. 

var. calcullensis Serv. 153 
var. malaccensis Serv. 153 
var. pallescens Serv. 156 
var. sepientrionalis Serv. 

var. silheiensis Serv. 153 
cumingii Schlechtend. 154 
ssp. perrollelii Serv. 154 
ssp. philippinensis Serv. 
dendroidea Sc\\\tz\\\sx\± 153 
ferruginea A. Rich. 153 
ssp. sumairana Serv. 153 
var. airovirens Serv. 153 
var. richardia Serv. 153 
fruticosa (Lour.) Cheval. 

gaudichaudiana Schlechtend. 

javanica Bl. 153 
latifoliaL. 152 
laiifolia (non L.) A. Rich. 
var. iriflora Schlechtend. 
/7erro»e/H Schlechtend. 154 
philippinensis Perrotlet 153 
rigida Bl. 153 
rosiraia Serv. 154 
iriilora Roxb. 152, 153 
ssp. obsoleia Serv. 153 
ssp. polymorpha Serv. 

ssp. rigida Serv. 153 

(Elaeagnus triflora) 

ssp. leiragonia Serv. 153 
ssp. leiragonia (non Serv.) 

Merr. & Perry 155 
var. brcvilimbata 't Han 

154*, 155 
var. brevilimhalus 155 
var. brevipes Serv. 153 
var. longipes Serv. 153 
var. triflora 154* 
Zollinger i Serv. 154 
Eiaeocarpus 123 
Elatinaceae 4: 203-206 
Elmerrillia Dandy 562, 564- 
567, 593, 595 
sect. P seudoaromadendron 

Dandy 595 
celebica (Koord.) Dandy 596 
mollis Dandy 596, 598 
ovalis (Miq.) Dandy 595, 

papuana (Schitr) Dandy 595, 
var. adpressa Dandy 596 
var. glaberrima Dandy 
596, 598 
platyphylla (Merr.) Noot. 

595, 596 
pubescens (Merr.) Dandy 595, 

sericea C.T.White 598 
tsiampacca (L.) Dandy 595, 
ssp. mollis (Dandy) Noot. 

567, 598, 599* 
ssp. tsiampacca 598 
var. glaberrima (Dandy) 

Noot. 595, 598 
var. tsiampacca 597*, 
vriesearuj (Miq.) Dandy 596 
Elsoia Adans. 483 

bracieala (Benn.) O.K. 484 
corymbosa (Turcz.) O.K. 

lavoyana (Benn.) O.K. 484 
Embelia urophylla Wall, ex 

A. DC. 649 
Emmenanihus Hk.f. & Am. 
622, 626 
chinensis Hk.f. & Am. 626 
Entosiphon Bedd. 665 
Epacridaceae 6: 422-444, 

943; 10: 335 
Ephedra 338 
Ephippiandra 258 
Epicryanlhes 489 
Epinetrum 164 
Epirhizanlhes 489 


Index to scientific plant names 


Epirixanthes Bl. 455, 456, 459, 
486, 488 
aphylla (Griff.) Merr. 490 
cylindricaBl. 457,490, 491 
elongata Bl. 489, 490* 
kinabaluensis Wendt 490, 

linearis Bl. 490 
pallida Wendt 490, 492 
papuana J.J. Sm. 489, 490, 

tenella Hk. f. 490 

Epirixanihus 489 

Epirizanthes 489 

Epirrhizanihe 489 

Epirrhizanlhes 489 

Eriandra Royen & Sleen. 455, 
457, 459, 492 
fragrans Royen & Sieen. 493 

Ericaceae 6: 469-914; 943; 
7: 827; 8: 549; 9: 562; 
10: 335, 716 

Ericybe Roxb. 29 

Eryihropalla 17 

Erythropalum Bl. 1-6, 17, 166 
grandifoliwn Elmer 17 
scandens Bl. 17, 18* 

var. abbrevialum Hochr. 17 
iriandrum Quis. & Merr. 29 
vagum (Griff.) Mast. 17 

Erythroropalum 17 

Erythroxylaceae 5: 543-552; 
8: 549; 10: 607, 609, 622 

Escalloniaceae 335 

Euglypha 53, 61,63 

Euphorbiaceae 29, 59 

Euiacta cunninghamii (Ait.) 
Link 426 

Eutassa Salisb. 425 

cunninghamii Spach 426 

Evodia 123 

Exellodendron Prance 642 
coriacca 64 1 

Exiielea Bl. 642 

Exiteles 673 


corymbosa (Bl.) Bl. 673 
muliiflora (Konh.) Walp. 

Eystathes Lour. 493, 507 

Fagaceae 7: 265-403; 8: 

549; 9: 563 
Faika Philipson 255. 262. 263. 

284 map 
villosa (Kanch. & Hatus.) 

Philip.son 284. 285*. 


Falcalifolium de Laub.343, 347, 
352, 354, 355, 371 
angustum de Laub. 372, 373 

map, 374 
falciforme (Pari.) de Laub. 

372, 373* map 
gruezoi de Laub. 372, 373 

papuanum de Laub. 372, 373 
Fawceltia F.v.M. 164, 188 
me rril liana (Diels) Yamamoto 
Ferolia O.K. 655 

asperula (Miq.) O.K. 671 
corymbosa (Bl.) O.K. 674 
cosiata (Konh.) O.K. 663 
glaberrima (Hassk.) O.K. 

griffithiana (Benth.) O.K. 

jackiana (Benth.) O.K. 671 
/jmda(Hk.f.)Ridl. 646 
nonda (F.v.M. ex Benth.) 

O.K. 658 
oblongifolia (Hk.f.)O.K. 

polyneura (Miq.) O.K. 664 
salicifolia (Presl) O.K. 674 
scabra (Hassk.) O.K. 669 
sumatrana (Jack) O.K. 661 
Fibraurea Lour. 157, 160-162, 
166-168, 170, 171, 207 
chloroleuca Miers 160,207 
chloroleuca (non Miers) Merr. 

elliplica Yamamoto 184 
fasciculata Miers 209 
haematocarpus Hk.f. & Th. 

laxa Miers 209 
recisa 207 

lincloria Lour. 160, 165, 
207, 208* 
Fibraureopsis Yamamoto 1 83 

smilacifolia Yamamoto 1 84 
Ficus/7u/cAra Wall. 323 
Fissipelalum Merr. 29 
Flacourlia camptoceras Miq. 

Flacourtiaceae 5: 1-106.565; 
6: 943; 7: 827; 9: 563; 
10: 333. 539, 715, 716 
Flagellariaceae 4: 245-250; 

5: 557; 9: 564 
Folium lunalum minus Rumph. 

Frcnclopsis 338 
Funis felleus Rumph. 194 

Ganua palembanica (Miq.) v.d. 
Assem & Kosterm. 453 

sp. 453 
Geobalanus Small 645 
Geraniaceae 6: 445-449; 

9: 565; 10: 607, 639 
Geraniales 609, 639 
Gesneriaceae 335 
Gigantopteris 338 
Gjellerupia Laut. 31-35, 45 

papuana Laut. 45*, 46 
Gladiolus dalenii Geel 717 

natalensis (Ecklon) Reinw. ex 
Hk. 717 
Glossocalyx 258 
Glyptopetalum loheri Merr. 716 
Gnetaceae 4: 336-347; 6: 

Goodeniaceae 5: 335-344; 
567; 6: 949; 7: 827; 9: 
Gordonia decandra Roxb. 626 
Govantesia Llanos 35 

malulucban Llanos 36 
Grossulariaceae 335 
Groutia Guill. & Perr. 46 

cellidifolia Guill. & Perr. 47 
Grymania Presl 67 1 

salicifolia Presl 671, 673 
Guatteria incerta Bl. 515 
Guttiferae 145 
Gymnostoma sp. 453 

Haematocarpus Miers 160, 161, 
167-169, 171, 183 
complus Miers 1 84 
incusus Miers 1 84 
subpeltatus Merr. 182*. 184 
ihomsonii Miers 184 
validus (Miers) Bakh.f. ex 
Forman 184 
Haemodoraceae 5: 111-113; 

10: 717 
Haemodorum coccineum R. Br. 
corymbosum Vahl 7 1 7 
Halocarpus 354 
Haloragaceae 7: 239-263, 

Ilamamelidaceae 5: 363- 

379; 6:952; 10: 341,717 
Harmandia Pierre ex Baill. 1-6. 
9. 10 map 
kunstleri King 9 
mckongensis Pierre ex Bail!. 
9, 10* 
Hcbcpctalum 608 
Ucckelia nymanii K.Sch. 253 


Flora Malesiana 

[ser. I, vol. 104 

Hedycarya 255, 259, 260 
arborea 256 

salomoncnsis Hemsl. 311 
Hedycrea Schrcb. 645 
Hcdyosmum Swarlz 123-129, 
143. 144 
arboresccns 125, 127 
brasiliense 143 
mexicanum 125 
nutans (non Sw.) Merr. 144 
orientale Merr. & Chun 

142*, 144 map 
sp. Sieen. 144 
sumatranum 144 
Hcisieria 3,4 
micranlha 3 
Hcnnecartia 257 
Ilernandia kunslleri 9 
Ueierosamara birmanica (O.K.) 

Chodat 463 
Heterotropa 53, 63 
Hexastylis 53, 63 
Hibbenia 123 
Himantandraceae 562 
Hippophae L. 151 
Ilolopeira Miers 231 
australis Miers 234 
laurifolia (DC.) Miers 234 
Holostylis 53,61,63 
Homalium dasyanthum 716 
Hopca gracilis Miq. 716 
Horsfieldia iryaghedi (Gaertn.) 

Warb. 605 
Hortonia 258, 259 
Hortoniaceae 261 
Hugonia L. 608, 609 
sect. Durandea (Planch.) 

BaUIon609, 611 
sect. Hugonia 611 
costata Miq. 610*, 611, 612 

jenkinsii F.v.M. 611, 612 

map, 613* 
pentagyna (Warb.) K.Sch. 

robinsonii Merr. 612 
sumatrana Miq. 618 
Hugoniaceae 607, 608, 629 
Humiria 629 

Humiriaceae 607, 609, 629, 630 
Hunga Pancher ex Prance 635- 
637, 642, 643, 650 
fusicarpa Kosterm. 647 
longifolia Prance 651, 653* 

novoguineensis Prance 651, 

652*, 653 map 
papuana (Baker f.) Prance 
651, 652*. 653 map 

Hyalisma 112 

Hydnocarpus nana King 716 
Hydrocaryaceae 4: 43-44 
Hydrocharitaceae 5: 381— 
413,569; 6: 952; 7: 
828; 9: 566; 10: 112, 
HydrophyUaceae 4: 207-209 
Hypericaceae 8: 1-29; 10: 

Hypericinea macrocarpa Wall. 

Hypericum henryi L6v. & Vaniot 
ssp. hancockii Robson 
uralum Buch.-Ham. ex 
D. Don 717 
Hyperixanthes 489 
Hypserpa Miers 157, 161, 167- 
169, 172, 222, 218 
borneensis (Miq.) Becc. 

cu5p/Jafa(Hk.f. &Th.) 
Miers 219 
var. imcrophylla (Miq.) 
Boerl. 219 
heteromera Miers 219 
jagorii'D'itXs 2\9 
latifolia Miq. ex Diels 221 
laurina (F.v.M.) Diels 218, 

monilifera (Burk.) Diels 

nandinifolia Yamamoto 

nitida Miers 219, 220* 
parvifolia Kaneh. & Hatus. 

polyandra Becc. 219, 221 
var. tomentosa Forman 
praevaricaia Miers 219 
propensa Miers 219 
raapii Diels 221 
selebica Becc. 221 
selwynii F.v.M. 219 
iriflora [non (DC.) Miers] 
Miers 219 
Hypsipodes Miq. 188 

Iberis amara L. 543 
umbellaia L. 543 
Icacinaceae 7: 1-87, 828; 9: 
566; 10: 29, 33, 91, 145, 
166, 629, 717 
Idenburgia Gibbs 145, 146, 
326, 333 
arfakensis Gibbs 149 


elaeocarpoides Gilg & Schllr 

novoguineensis Gibbs 147 
pachyphylla Gilg & Schltr 

pauciflora A.C. Smith 147 
Indorouchera HaU.f. 607-609, 
contestiana (Pierre) Hall. f. 

616*, 619 map 
griffithiana (Planch.) Hall.f. 

616*, 617*, 618* map 
rhamnifoUaWaXU. 6\9 
Iridaceae 8: 77-84; 10: 717 
Irina integer rima Bl. 702 
Irvingiaceae 621 , 622 
Isomerocarpa A.C. Smith 263 
novoguineensis (Perkins) 
A.C. Smith p.p. 265, 266 
Ixionanthes 622 
Ixonanthaceae 607, 609, 621- 

627, 629 
Ixonanthes Jack 621, 622, 623* 
sect. Brewstera (M.J. 

Roemer) Hall.f. 621, 625 
sect. Emmenanthes Hall.f. 

sect. Ixonanthes 621, 625, 

chinensis (Hk. f. & Am.) 

Champ. 626 
cochinchinensis Pierre 626 
crassifolia Hall.f. 627 
cuneata Miq. 625 
dodecandra Griff. 625 
grandiflora Hochr. 626 
grandifolia Ridl. 627 
hancei Pierre 626 
icosandra Jack 624*, 625 
var. cuneata Miq. 625 
var. obovata Ridl. 625 
khasiana Hk. f. 626 
longipedunculata Merr. 

lucida Bl. 625 
multiflora Stapf ex Ridl. 

obovata mi.i. 615 
papuana (Schltr) Hub.Winkler 

petiolaris Bl. 623, 625, 626, 

petiolaris (non Bl.) Hall.f. 

philippinensis Elmer 626 
reticulata Jack 625, 626 
subdodecandra 625 


Index to scientific plant names 


Jackia longifolia 514 

vitellina 514 
Jacquemontia browniana Oosistr. 
pannosa (R.Br.) Mabberley 
Jakkia Bl. 493, 508 
longifolia Bl 514 
vitellina Bl. 514 
Juglandaceae 6: 143-154, 

Juncaceae 4: 210-215; 5: 

557; 6: 953; 9: 566 
Juncaginaceae 4: 57; 5: 554 
Juniperus 345 
chinensis 453 
elata Roxb. 363 
Juppia borneensis Merr. 253 

Kairoa Philipson 255, 256, 258, 

259, 262, 263, 284 map, 

suberosa Philipson 304*, 
305, 306* 
Kibara Endl. 255, 257, 259, 

260, 262, 263, 287, 308 
angusiifolia Perkins 298 
archboldiana A.C. Smith 

257, 289, 291, 293 
arucnsis Becc. 305 
blumei Sleud. 298 
borneensis Boerl. 305 
buergersiana Perkins 305 
bullata Philipson 289, 

carrii Philipson 257, 290, 

291, 297 
chariacea Bl. 298 
chimbuensis Philipson 290, 

clemensiae Perkins 298 
coriacea (Bl.) Tulasne 262, 

290, 298 
cuspidaia Bl. 298 
depauperata Merr. 3(X) 
dichasialis Suesscng. & 

Heme 298 
ellipsoidea Merr. 298 
clmeri Perkins 305 
clongata A.C. Smilh 290. 

fcrox Philipson 257. 258. 

289. 291. 292* 
flagclliformis Philipson 290. 

form icar urn Becc. 305 
fragrans Philipson 290. 300 


fugax Philipson 290, 300, 

301, 302* 
grandifolia Merr. 298 
hartleyi Philipson 289, 295, 

hirsutaV^aih. 315 
hospitans Becc. 311 
inamoena Perkins 298 
karengana Philipson 289, 

katikii Philipson 290, 298 
kosiermansii Philipson 289, 

lalifolia Philipson 257, 289, 

laurifolia A.C. Smilh 290. 

leachii Philipson 289, 294 
ledermannii Perkins 296 
longipes Perkins 298 
macrantha Philipson 288*. 

289, 297, 300 
macrocarpa Perkins 298 
macrophylla Perkins 298 
merrilliana Perkins 298 
microphylla Perkins 289, 

mollis Merr. 298 
moluccana Perkins 288*, 

289, 295, 296, 303 
monticolia Perkins 290, 303 
motleyi Perkins 298 
myrtoidea Perkins 289, 291 
ncriifolia Perkins 305 
nitens Philipson 290, 299 
novobrilanica Philipson 290, 

oblongata Philipson 288*. 

289, 295 
oblusa Bl. 290, 300 
oligocarpclla (Kaneh. & 

Hatus.) Philipson 256, 

289, 291 
olivaeformis Becc. 305 
papuana A.C. Smith 290. 

pcrkinsiae K.Sch. & Laul. 

rigidifolia A.C. Smilh 256. 

258, 288, 290, 302 
rocmeri (Perkins) Philip.son 

289, 290 
rossclcnsis Philipson 289. 

roycnii Philipson 289, 295 
serrulula Pcrkms 298 
shungolcnsis Philipson 289. 



sleumeri Philipson 290, 303 
stapfiana Perkins 298 
streimannii Philipson 288, 

289, 296 
sudestensis Philipson 290, 

symplocoides Perkins 289, 

290, 303 
leijsmanniana Perkins 296 
timorensis Boerl. 305 
tomeniosa Perkins 298 
irichantha Perkins 298 
versieeghii Philipson 289, 

vidalii Perkins 298 
vrieseana Perkins 296 
warburgii Perkins 298 
warenensis Kaneh. & Hatus. 
289, 295 
Kibaropsis 260 
Kingsboroughia Liebm. 690 
Kmeria 566 
Korihalsella 353 

dacrydii 353 
Kosiermanthus I*rance 636- 
639, 641, 642, 675 
heteropetalus (Scort. ex King) 
Prance 676*. 677 map, 
malayanus (Kosterm.) Prance 
677 map 
Krameriaceae 457, 458 

Labiatae 8: 301-394; 9: 

Lagarosirohus Quinn 360, 361 
Lardizabalaceae 163, 679 
Larix 343 

Lauralcs 127,327,330 
Lautcrbachia Perkins 263. 284 
map, 326 
novoguineensis Perkins 
325*. 326 
Lavallea Baill. 19 

ceylanica (Gardn.) Baill. 

philippinensis Baill. 22 
Leeaceae 7: 755-782 
Legnephora Micrs 158. 165, 
169, 170, 172, 225 
acuta Forman 159. 225. 

microcarpa Forman 225. 

226*. 227 
minutidora (K.Sch.) Dicis 
225. 226*. 227. 228* 


Flora Malesiana 

[ser. I, vol. 104 


moorei (F.v.M.) Miers 225, 

nyctericarpa Diels 227 
philippinensis Forman 225, 
Lemnaceae 7: 219-237 
Lentibulariaceae 8: 275-300 
Lepidium L. 541,543,545,547 
lacteviride (P. Royen) 
Hewson 548, 549* 
maccowagei Hews. 547, 

minuliflorum (Ridl.) Hews. 

sativum L. 547, 548 
virginicum L. 547, 548 
Lepidocarpa Korlh. 655 
costata Korth. 663 
ovalis (Korth.) Bl. ex Miq. 
Lepidocarya costata 663 
Lepidothamnus Phil. 360 
Lepionurus Bl. 31-35, 42, 43 
javanicus G. Don 43 
oblongifolius (Griff.) Mast. 
var. angustifoUus Roxb. 
pubescens Ridl. 52 
sylvestris Bl. 31, 33, 34, 43, 
44* map, 46 
var. lanceolata Val. 43 
Leptonium Griff. 43 

oblongifolium Griff. 43 
Levieria Becc. 263, 275 
acuminata (F.v.M.) Perkins 

277, 279 
beccariana Perkins 277, 280*, 

forbesii Perkins 281, 282 
laxiflora Perkins 279, 281 
montana Becc. 277, 279, 

montana (non Becc.) Kaneh. 

& Hatus. 277 
nitens Perkins 277, 278 
orientalis Philipson 277, 278 
parvifolia A.C. Smith 277 
rudolfii Perkins 278 
scandens Philipson 277, 279 
schlechteri Perkins 278, 279, 

squarrosa Perkins 276*, 277, 

urophylla Perkins 279, 281 
Levieria Kosterm. p.p. 263 
Libocedrus Endl. 337, 339, 340, 
342, 347, 443, 444 map 


subg. Eulibocedrus Pilger 

arfakensis Gibbs 446 
papuana F.v.M. 342, 444, 

var. arfakensis (Gibbs) 
de Laub. 446, 447 map 

var. papuana 445, 446*, 
447 map 
torricellensis Schltr ex Laut. 

Licania Aubl. 635-640, 642, 

643, 645 
angelesia Bl. 646 
diemenia Bl. 649 
elaeosperma 640 
fusicarpa (Kosterm.) Prance 

646, 647, 650 map 
macrophylla 639 
palawanensis Prance 646, 

647 map 
splendens (Korth.) Prance 

640, 646, 647 map 
Liliaceae 9: 189-235; 10: 

109, 112, 166 
Limacia Lour. 158, 160, 161, 

167, 168, 170, 172, 221, 

blumei (Boerl.) Diels 222, 

borneensis Miq. 219 
cerasifera Becc. 224 
cerasifolia 224 
cuspidata Ek.f.&Th. 219 
distincta Miers 223 
inornata Miers 223 
kunstleri King 233 
microphylla Miq. 219 
monilifera Burk. 221 
nativitatisRidl. 218 
oblongaHk.f. & Th. 160, 

165, 166, 223 
scandens Lour. 220*, 223 
selwynii (F.v.M.) Bailey 

sumatrana Scheff. 198 
triandra (Colebr.) Hk.f. & Th. 

velutina Hk.f. & Th. 160, 

var. glabrescens King 223 
Limnanthaceae 639 
Linaceae 29, 607-619, 621, 

622, 629, 630, 639 
subfam. Hugonioideae 607, 

608, 609 
subfam. Ixonanthoideae 621 
subfam. Linoideae 607, 609 

Linum 609 

Liriodendron 562, 563, 565- 
567, 605 
liliiferum L. 582 
tulipifera 567 
Liriopsis Spach 598 

fuscata (Andr.) Spach 598 
Litsea baticulin (Blco) Kosterm. 
insignis (Bl.) Boerl. 539 
leytensis Merr. 717 
Lobbia Planch. 65 

de pendens Planch. 75 
Lobularia maritima (L.) Desv. 

Loganiaceae 6: 293-387, 

953,960; 7: 828; 9:567 
Longpistillate Manglieiia Meijer 

Lophopetalum beccarianum 

Pierre 716 
Lophopyxidaceae 7: 89-91 
Lophostylis Hochst. 483 

javanica Miq. 484 
Loranthaceae 2, 3, 34, 35 

Macadamia 123 

Macrococculus Becc. 157, 158, 
160-162, 167-169, 171, 
pomiferus Becc. 159, 178, 

179, 180* 
tympanopodus Laut. & 
K.Sch. 211 
Madhuca burckiana (Koord.) 

Lamk 593 
Magnolia L. 564, 566, 567, 
sect. Gwillimia 566, 569 
sect. Lirianthe 565 
sect. Liriopsis Baillon 598 
sect. Maingola 565, 567 
sect. Manglietia (Bl.) Baillon 

sect. Talauma Baillon 568 
sect. Theorodon 569 
subg. Magnolia 564-567, 
sect. Maingola Dandy 569, 
subg. Talauma (Juss.) Pierre 
564-567, 576 
sect. Aromadendron (Bl.) 
Noot. 566, 569, 576 
sect. Blumiana Bl. 565, 
566, 569, 581 
subg. Talauma Pierre 568 
subg. Yulania 565 


Index to scientific plant names 



aeqidnoctialis Dandy 57 1 
angaiensis Blco 585 
ashtonii Dandy ex Ncx)t. 570, 

576, 578*, 579 
ashtonii Noot. 579 
beccariana (Agoslini) Nc»L 

betongensis (Craib) H. Keng. 

bintuluensis (Agoslini) Noot. 

570, 576, 577 
blumei Pranil 589 
bomeensis Noot. 570, 576, 

579, 580* 
candollii (Bl.) H. Keng 569, 

581, 582 

var. angaiensis (Blco) 

Noot. 582, 585 
var. beccarii (Ridl.) Nool. 

582, 586 

var. candollii 582, 584* 

var. obovaia (Korih.) 
Nooi. 582, 585 

var. singapurensis (Ridl.) 
Nool. 582, 586 
carsonii Dandy ex Nool. 570, 

var. carsonii 572*, 573, 

var. drymifolia Nool. 
573, 574* 
carsonii Dandy ex Cockbum 

571, 572 
coco (Lour.) DC. 569 
decandollti 5S\, 5^2 
denudata 562 
drymifolia Dandy ex 

Cockbum 571, 573 
elegans (HI.) H. Keng 570. 

576. 577 
forbesii King 582 
fragrans Reinw. ex Bl. 

giganlifolia (Miq.) Nool. 

581, 586 
grandi flora L. 569 
javanica K. & V. 571 
kachirachirai 563 
kunsllen King 582 
lasia Noot. 581. 587 
lilufera (L.) Baillon 582 
macklottii (Korlh.) Dandy 
569. 570. 571 

var. bctcariana (Agoslini) 
N(x)l. 571 

var. macklottii 571 
maingayi King 570. 571, 



maingayi (non King) Ridl. 

mariusjacobsia NooL 581. 

nilida 563 

nutans (Dandy) H. Keng 577 
odoratissima Reinw. ex Bl. 

pachyphylla Dandy 583 
paenetalauma 566 
pahangensis Noot. 570. 576, 

pealiana (non King) K. & V. 

pealii 571 

persuaveolens Dandy 581, 
ssp. persuaveolens 587 
ssp. rigida Nool. 587 
var. pubescens NooL 

587. 588 

var. rigida 587 

phaulantha Dandy ex Noot. 

570. 571. 576 
Philippine nsis Parmeni. 604 
plumierii Schwartz 568 
pumila aucl. 582. 583 
rumphii Spr. 582 
sarawakensis (Agoslini) 

Noot. 581, 588 
singapurensis (Ridl.) H. Keng 

splendens Reinw. ex Bl. 582 
sprengeri 567 
stellata 562 
uvariafolia Dandy ex 

Cockbum 574 
uvariifolia Dandy ex Noot. 

570. 571. 574, 575* 
villosa (Miq.) H. Keng 581. 

virginiana L. 568 
vrieseana (Miq.) Baillon ex 
Pierre 596 
Magnoliaceae 261. 561-605 
subfam. Liriodendroidcac 561. 

subfam. Magnolioideae 561. 
563. 564. 567. 568 
tribe Magnolieae 561, 

tribe Michelicac Law Yuh- 
wu 561. 568. 593 
Magnolialcs 62, 64, 261 
Maingaya malayana Oliver 7 1 7 
Malania 1 

Maipighiaceae 5: 125-145, 
566; 6: 960; 10: 458 

Malulucban 35 
Malvanae 151 

Manglietia Bl. 564-568, 589 
calophylla Dandy 589, 591 
candollii (Bl.) WaU. 582 
celebica Miq. 582 
dolichogyna Dandy ex NooL 

589, 590*, 591 
dolichogyna Dandy ex 

Cockbum 591 
glauca Bl. 589, 590* 
var. glauca 589, 590 
var. lanuginosa Dandy 

var. sumatrana Dandy 
589, 591 
glauca (non Bl.) King 591 
g/auca (non Bl.) Ridl. 591 
glauca aucL 585 
lanuginosa (Dandy) Nool. 

589, 591 
macklottii Korlh. 571 
macklottii aucl. 589 
minahassae K. & V. 593 
oortii Korlh. 577 
oortii (non Korlh.) Miq. 577, 

oortii aucL 589 
pilosa Parmeni. 590. 591 
sabahensis Dandy ex Nool. 

589. 591. 592* 
sabahensis Dandy ex 
Cockbum 591 
scortechinii King 60 1 
sebassa King 582 
singalanensis Agoslini 590 
sumatrana Miq. 589, 591 
Manglieliastrum 563 
Martyniaceae, see Pedaliaccac 
Malthaea Bl. 255, 258, 262, 
263, 288, 319, 321 map 
calophylla Perkins 323 
charlacea Merr. 321 map, 

322*. 323 
heierophylla Quis. & Merr. 

321 map, 322*. 323.324 
intermedia Merr. 321 map, 

322*, 323, 324 
laiifolia Perkins 323 
philippinensis Perkins 326 
pincholiana Perkins 323 
pubescens Merr. ex Perkins 

321 map. 322*. 323 
roemcri Perkins 290 
sancta Bl. 320*. 321 map. 
322*. 323 
var. mindanaoensis 

Perkins 323 
var. venulosa Perkins 323 


Flora Malesiana 

[ser. I, vol. 104 


vidalii Perkins 321 map, 

322*. 323, 324 
williamsii Perkins 323 
Matlhiola incana (L.) R.Br. 545 
Meiogyne virgata (BI.) Miq. 716 
Melientha Pierre 31-34, 38, 40 
acuminata Merr. 39 
suavis Pierre 34, 39* 
ssp. macrocarpa Hiepko 

40 map 
ssp. suavis 40* map 
Meliosma Bl. 163, 679-682, 
690, 691*, 692* 
subg. Kingsboroughia 

(Liebm.)Beus. 691*. 
sect. Hendersonia Beus. 

692*, 694 
sect. Kingsboroughia 
692*, 694 
subg. Meliosma 691*, 693, 
sect. Lorenzanea (Liebm.) 

Beus. 694 
sect. Meliosma 692*, 693 
subsect. Pinnatae 
(Warb.) Beus. 693, 
subsect. Simplices 
(Warb.) Beus. 693, 

ser. Curvinervia 

Beus. 693 
ser. Rectinervia 
Beus. 693 
acutrdnatissima Merr. 712 
angulata Bl. 690, 698 
apoensis Elmer 709 
arnoitiana Walp. 709 
bartlettUM&n. 710 
bonlocensis Merr. 697 
brachyboirys Merr. 713 
cannarioides Elmer 709 
celebica Warb. ex Dihm 

conferiiflora Merr. & Perry 

confusa Bl. 702 

var. laxior Baker f. 713 
costata CvSod. 701 
cuspidata Bl. 702 
diepenhorstii Valet. 702 
dolichomischa Vidal 696 
elegans Rm. 110 
ellipticaHk.f. 701 
elmeri Merr. 702 
evrardii Gagnep. 701 


ferruginea Bl. 711 
ferruginea (non Bl.) Backer & 

Bakh.f. 709 
ferruginea (non Bl.) Koord. 

ferruginea (non Bl.) Merr. & 

Perry 711 
floribunda Bl. 709 
fruticosa Bl. 701 
glauca Bl. 709 

var. floribunda (Bl.) 
K. & V. 709 
glomerulaia Rchd. & Wils. 

grandifolia Lecomte 713 
harmandiana Pierre 699 
hirsuta Bl. 694, 703 map, 

705, 715 
humilis Merr. & Perry 

lanceolata Bl. 694, 703 map, 
704, 715 
var. chariacea Bl. 704 
var. genuina Hochr. 704 
var. lanceolata 705 
f. lanceolata 705 
f. nervosa (K. & V.) 
Beus. 694, 705 
var. membranacea Bl. 704 
var. obliqua Bl. 704 
var. obliqua (non Bl.) 

Koord. 711 
var. pendula Bl. 704 
var. polyptera (Miq.) Beus. 
703 map, 705 
/anc//b//aHk.f. 701 
laurina BIT \5 
lepidota Bl. 694, 695 

ssp. dolichomischa (Vidal) 

Beus. 695, 696* 
ssp. kinabaluensis Beus. 

695, 697 
ssp. lepidota 695 
ssp. squamulata (Hance) 

Beus. 697 
ssp. vulcanica (Merr.) 
Beus. 695, 697 
levis King 704 
loheri Merr. 701 
luzonensis l(y) 
luzonica 709 
macgregorii Merr. 712 
macrocarpa Elmer 7 1 1 
macrophylla Merr. 711 
megaloboirys Merr. 711 
monophylla Merr. 701 
monophylla Ridl. 696 

multiflora 709 

myriantha Sieb. & Zucc. 681 
nervosa K. & V. 704, 705 
nitida Bl. 702 

var. cerasiformis Bl. 702 
var. splendens BI. 702 
var. tridenta Bl. 702 
pannosa Hand.-Mazz. 701 
patens Hemsl. ex Forb. & 

Hemsl. 699 
paucinervia Merr. 710 
pedicellata K. & V. 695 
pedicellata (non K. & V.) 

Merr. 697 
pendula Merr. 712 
petiolaris Miq. 715 
philippinensis Merr. & Perry 

pinnata (Roxb.) Maxim. 694, 
ssp. amottiana (Wight) 
Beus. 707, 708 map, 
ssp. ferruginea (Bl.) Beus. 
707, 708 map, 709, 
ssp. humilis (Merr. & 
Perry) Beus. 707, 708 
map, 713 
ssp. macrophylla (Merr.) 
Beus. 706*, 707, 708 
map, 709, 711 
ssp. pendula (Merr.) Beus. 
707, 708 map, 709, 
ssp. ridleyi (King) Beus. 
707, 708 map, 709, 
ssp. sylvatica (Elmer) 
Beus. 707, 708 map, 
709, 712 
pinnata Koord. 702 
polyptera Miq. 704, 705 
pungens (Wall, ex W. & A.) 

Walp. 702 
pungens auct. 699 
reticulata Merr. 712 
ridleyi King 710 
rigida Sieb. & Zucc. 699 
var. angustifolia Miq. 

var. patens Cufod. 699 
rufo-pilosa Hend. 694, 703 

map, 714 
sambucina Miq. 709 
sarawakensis 694 
schlechteri Merr. & Perry 


Index to scientific plant names 



simplicifolia (Roxb.) Walp. 
681, 694, 697*, 698, 
699 map 
ssp. fruticosa (Bl.) Beus. 
697*, 698, 699 map, 
700*, 701 
ssp. pungens (Wall, ex 
W. & A.) Beus. 697*, 
698, 699*. 702 
ssp. rigida (Sieb. & Zucc.) 
Beus. 697*, 698, 699 
ssp. simplicifolia 690, 
697*, 698, 699 map 
sumatrana (Jack) Walp. 694, 

702, 703 map, 715 
sylvatica Elmer 712 
limorensis Bl. ex Blenk 715 
tongcalingii Elmer 711 
trichocarpa Merr. 710 
vulcanica Mcrr. 697 
wallichii (non Planch, ex 

Hk.f.)Koord. 711 
wightii Planch. 702 
Meliosmaceae End!. 679, 680 
Memecylanthus Gilg & Schltr 

Meniscosta Bl. 682 
javanica Bl. 684 
scandens Bl. ex Spr. 684 
Menispermaceae 19, 157- 
253, 305, 679, 680. 
iribc Coscinieac 161, 162, 

iribe Fibraureeae 162, 164, 

tribe Menispermeae 161, 162, 

164, 168 
iribc Tiliacoreae 162, 164, 

iribc Tinosporcae 158, 161, 
162, 164, 167, 168, 189* 
iribc Triclisieae 162 
Mcnispcrmum 158, 165 
cocculus L. 213 
crispum L. 194. 195 
dauhcum 165 
fenesiralum Gaerin. 215 
flavescens Lamk 210 
flavum L. 210 
glabrumBumn.f. 199,200 
glaucum Lamk 229 
japonicum TTiunb. 245 
lacunosum Lamk 213 
limacia (DC.) Sprcng. 

nuUabaricum Lamk 200 


orbiculatum L. 231 
ovalifolium Vahl ex Pers. 
iriandrumRoxh. 185 
trilobum Thunb. 231 
tuber culalum Lamk 194 
Metasequoia 345 
Michelia L. 562, 564-567. 

593, 595, 598, 605 
X alba DC. 601,605 
arfakiana Agoslini 596 
beccariana Agoslini 571 
blumei Steud. 601 
celebica Koord. 596 
champaca L. 562, 565, 598, 

var. champaca 603 

var. pubinervia (Bl.) Miq. 
champaca Miq. 603 
champacca aucl. 596 
cumingii Merr. 604 
dollsopa auct. 589 
ecicatrisaia Miq. 603 
figo (Lour.) Spr. 599, 605 
forbesii Baker f. 596 
fuscala 562 
koordcrsiana Nooi. 601, 

longiflora 605 
longifolia Bl. 605 

var. racemosa Bl. 605 
mollis (Dandy) McLauchlin 

moniana Bl. 563, 601, 603, 
604 map 

var. subvelulina Miq. 604 
montana aucl. 603 
parviflora 604 
philippinensis (Parment.) 

Dandy 601,604 
p////era Bakh.f. 603 
platyphylla Merr. 596 
pubinervia Bl. 603 
rufinervis Bl. 603 
salicifolia Agoslini 601, 604 
scoriechinii (King) Dandy 

600*, 601 
spec. Slapf 587 
suaveolens Pers. 601 
sumatrae Dandy 604 
isiampacca Bl. 603 
tsiampacca L. 596 

var. blumei Mor. 603 
velutina Bl. 603 
Microcachrys338. 343. 351. 

Microcarpus 35) 
Microdcsmis Hk.f. 29 

Microtinomiscium 158 
Milling tonia Roxb. 690 
arnoitiana Wighl 709 
ferruginea Schull. & Schull. 

lanceolata Schull. & Schull. 

nitida Schull. & Schull. 702 
pinnata Roxb. 707 
pungens Wall, ex W. & A. 

sambucina Jungh. 709 
simplicifolia Roxb. 697, 698 
sumatrana Jack 702 
Minquarlia 3 
Miquelia 166 
Mirtana loureiri (Pierre) Pierre 

Modeccopsis vaga Griff. 17 
Mollinedia 256 

acuminata F.v.M. 279 
coriacea (Bl.) Baill. 298 
sancta (Bl.) Baill. 323 
Molluginaceae, sec Aizoaceae 
Monaria Konh. ex Val. 17 
Monimia 257-261 

oval i folia 257 
Monimiaceae 145,255-326, 
256 map, 327, 330 
subfam. AUierospcrmaloidcac 

255, 258-263 
subfam. Horionioideae 260, 

subfam. Mollinedioideae 259, 

subfam. Monimioideae 259- 

263, 277 
subfam. Siparunoideae 258- 
tribe Hcdycaryeae 255, 259, 

262, 263, 277 

tribe Mollincdieae 255, 262, 

263, 277, 284 map, 326 
tribe Monimieae 259 
tribe Trimcnicac 327 

Monnina 456 
ejcce/.va(BI.)Spr. 501 
longifolia {E\.)S\)v.5\A 
macrophylla Slcud. 514 
vj/e/Z/na (Bl.) Spr. 514 
Monoon incertum (Bl.) Miq. 

Monquarlia 4 
Moquilea Aubl. 645 

sect. Cyclandrophora (Ha.ssk.) 
Endl. 665 
Morinjjaccae 4: 45—46; 5: 
554; 6: 960 


Flora Malesiana 

[ser. I, vol. 104 

Moutabea Aubl. 456, 493 
Munnickia Bl. ex Rchb. 65 
Myoporaceae 4: 265—266; 

9: 568 
Myricaceae 4: 276-279 
Myristicaceae 605 
Myrtales 151 
Myrtanae 1 5 1 
Myrtiflorae 151 

Nageia Gaertn. 337, 343, 347, 

353-355, 389, 391 map 
sect. Nageia 390 
sect. Polypodiopsis (Bertrand) 

de Laub. 390, 394 
amara (Bl.) O.K. 387 
beccarii (Pari.) Gordon 393 
blumei (Endl.) Gordon 393 
cmot//i^/7 (Pari.) O.K. 381 
discolor (Bl.) O.K. 400 
falciformis (Pari.) O.K. 372 
leptosiachya (Bl.) O.K. 400 
maximus (de Laub.) de Laub. 

339, 394 map 
motleyi (Pari.) de Laub. 393, 

394 map 
neglecta (Bl.) O.K. 400 
neriifolia (D. Don) O.K. 400 
polystachyus (R. Br. ex 

Endl.) O.K. 417 
/■Mm/7;://(Bl.)F.v.M. 415 
teysmannii (Miq.) O.K. 406 
thevetiaefolia (Bl.) F.v.M. 

vitiensis (Seem.) O.K. 394 

wallichiana (Presl) O.K. 

391, 392*, 393 map 
Najadaceae 6: 157—171 
Nallogia Baill. 35 

gaudichaudiana Baill. 36 
Nasturtium R.Br. 555 
backeri Schulz 556 
benghalense DC. 560 
diffusum auct. 559 
heterophyllum Bl. 559 
homalospermum Schulz 557 

var. macrocarpum Schulz 
hybospermum Schulz 559 
indicum (L.) DC. 

vai.javana Bl. 559 
indicum auct. 557, 559 
novo-guineense Gilli 557 
officinale R.Bt. 555 
palustre (L.) DC. 557 
peekelii Schulz 557 
schlechieri Schulz 559 

Natsiatum Rheede 213 
Ncmuaron 256 
Neocarya Prance 642 

macrophylla (Sabine) Prance 
Nephroia Lour. 231 
elegans Ridl. 233 
sarmentosa Lour. 233 
Nigrina Thunb. 129 
spicala Thunb. 133 
spicifera Lamk 133 
Nouhuysia Laut. 145, 146 

arfakensis (Gibbs) Steen. 149 
novoguineensis (Gibbs) 

Hatus. 147 
pachyphylla (Gilg & Schltr) 

Hatus. 147 
papuana Laut. 147 
pauciflora (A.C. Smitji) 
Steen. 147 
Nyctaginaceae 6: 450-468; 

7: 829 
Nyssaceae 4: 29-31 

Ochanostachys Mast. 1-6, 12, 
14 map 

amentacea Mast. 5, 13*, 14 

bancana (Becc.) Val. 14 
Ochnaceae 7: 97-119; 10: 

145, 621 
Ochthocosmus Benth. 621 
Octarillumfruticosum Lour. 

Octoknema 3 
Octoknemaceae 33 
Odontocarya 158, 164 
Odonlocaryioides 158 
Olacaceae 1-29, 166, 629, 

tribe Agonandreae 34 

tribe Anacoloseae 3 

tribe Aptandreae 4 

tribe Couleae 4 

tribe Heisterieae 4 

tribe Olaceae 4 

tribe Opilieae 34 

tribe Schoepfieae 4 
Olacales 33, 34 
Olacinea ignota 25 
Olacineae 29, 34 
Olax L. 1-5,6 

sect. Triandrae Engl. 7 

baticulin Blco717 

benthamiana 3 

imbricata Roxb. 7, 8 

laxiflora Ridl. 9 

muliiflora A. Rich, ex Baill. 8 

multiflora Ridl. 9 


obtusa Bl. 7 
psittacorum (Willd.) Vahl 

rosea Ridl. 9 
scandens Roxb. 2, 7, 8*, 9, 

semiinfera Val. 9 
stricta 2 

sumatrana Miq. 9, 50 
Onagraceae 8: 98-113 
Ongokea 3, 4 
Ophiocaryon Schomb. 163, 679, 

Opilia Roxb. 4, 31-35,46 
sect. Lepionurus (Bl.) Baill. 
sect. Opiliaslrum Baill. 35 
subg. Urobotrya (StapQ Engl. 

acuminata Wall. 44 
amentacea Roxb. 31-34, 47 

celtidifolia (Guill.& Perr.) 
Endl. ex Walp. 33, 47 
cumingiana Baill. 36 
fragrans Elmer 47 
javanica Miq. 47 
manillana Baill. 36 
pentitdis Bl. 47 
thorelii Gagn. 47 
tomentella (Oliv.) Engl. 47 
Opiliaceae 2, 9, 29, 31-52 

tribe Opilieae 34 
Osmoxylon sessiliflorum (Laut.) 

Philipson 716 
Oxalidaceae 7: 151-178, 
829; 10: 607 

Pachydiscus Gilg & Schltr 335 
Pachygone Miers 160, 165, 167, 
168, 170, 172, 217 
brachystachya (DC.) Miers 

hebephylla Miers 217, 218 
leplostachya (DC.) Miers 

ovata (Poir.) Hk.f. & Th. 
217, 218, 220* 
var. dasyphylla Miq. 217 
var. rolundifolia Miq. 
pubescens Benth. 217, 218 
zeylanica Sant. & Wagh 
Pachylamax Dandy 561, 567, 
568, 593 
praecalva Dandy 593, 594* 
Paelae 493 


Index to scientific plant names 


Palmeria F.v.M. 255, 257-263, 

acuminata Kaneh. & Halus. 

270, 271 
angica Kaneh. & Hatus. 267, 

arfakiana Becc. 262, 267, 

brassii Philipson 269, 271, 

clemensae Philipson 268*, 

269, 273 
dallmannensis Kaneh. & 

Hatus. 272 
fengeriana Perkins 269, 271 
gracilis Perkins 267, 269, 

habbemensis A.C. Smith 272 
hooglandii Philipson 267, 

hypargyrea Perkins 269, 272 
hypochrysea Perkins 275 
incana A.C. Smith 269, 272 
montana A.C. Smith 269, 

272, 275 
myrianiha Perkins 270, 271 
myrtifoUa Perkins 270 
paniculate Ridl. 269 
parvifolia Kaneh. & Halus. 

puberula A.C. Smith 270 
pulchra Perkins 270, 271 
pulteana Perkins 272 
scandens 272 

schoddei Philipson 269, 275 
warburgii Perkins 270, 27 1 
womersleyi Philipson 269, 

273, 274* 
Pandaccae 29 

Papaveraceae 5: 114-117 
Papuacedrus Li 443 

arfakensis (Gibbs) Li 446 
papuana (F.v.M.) Li 444 
torricellensis (Schllr) Li 445 
/'apuz/V/a Ridl. 541,547 
laeteviridis P. Royen 548 
mmutiflora Ridl. 548 
minutiflora auct. 549 
Parabacna Micrs 158, 161, 162, 
167, 168, 170. 171, 201 
amplifolia Diels 193 
cincinnans (K. Sch.) Diels 

dcnudata Diels 189*. 202. 
. cchinocarpa Diels 189*. 201. 
202, 204 
var. pubescens Yamamoto 


elmeri Diels 189*, 202 

var. philippinensis 

(Merr.) Yamamoto 203 
hirsuta (Becc.) Diels 193 
hirsuta (non Becc.) Diels 202 
megalocarpa Merr. 158, 

186*, 189*. 202 
myriaditha 203 
myrianiha K. Sch. 203 
philippinensis Merr. 203 
sagiitaia Miers 189*, 201, 

scylophylla Diels 197 
tuberculata Becc. 189*, 202, 

Paracryphiaceae 145 
Parakibara Philipson 255, 262, 

263, 284 map, 286 
clavigera Philipson 286, 

Paramanglielia Hu & Cheng 

aromatica (Dandy) Hu & 

Cheng 589 
Paramichelia H.H. Hu 565, 567, 

593, 599 
baillonii (Pierre) Hu 599 
scoriechinii (King) Dandy 

Pararistolochia 53, 57, 59, 61, 

Parasitaxus 343, 351, 354 
Paraslemon A. DC. 635-638, 

640, 642, 648 
grandifructus Prance 649, 

650 map 
spicalus Ridl. 649 
urophyllus (Wall, ex A. DC.) 

A. DC. 637, 648*. 649, 

650 map 
versteeghii Merr. & Perry 

637, 649, 650 map 
Parinari Aubl. 635, 636, 639- 

642, 654 
seel. Exitelia (Bl.) C. Mucll. 

sect. Neocarya DC. 64 1 
sect. Pclrocarya 64 1 
sect. Sarcoslegia Benth. 671 
subg. Euparinari 654 
subg. Exitelia Bl. 673 
subg. Sarcostegia (Bcnlh.) 

Miq. 671 
anamensis 656 
argcnlco-sericea Kosierm. 

655, 656 map 
ashtonii Koslcrm. 660 
asperula auci. 667 


bicolor Merr. 664 
campesu-is 641 
canarioides Kosterm. 636, 

655, 656 map 
coriacea Benth. 641 
costata (Korth.) Bl. 636, 656, 
ssp. costata 663, 664 map 
ssp. polyneura (Miq.) 
Prance 655, 663, 664 
ssp. rubiginosa (Ridl.) 
Prance 663, 664 map 
elmeri Merr. 655, 657 map 
excelsa Sabine 639, 641 
giganiea Kosterm. 636, 655, 

660 map 
glaberrima Hassk. 641 
glaberrimiun (Hassk.) Hassk. 


var. lanceolatum (T. & B.) 
K. & V. 669 
griffithiana 64 1 
heteropetala Scort. ex King 

insularum 639, 656 
jackiana Benth. 641, 671 
latifrons Kosterm. 666 
macrophylla Sabine 641 
meiallica Kosterm. 656, 660 

montana 641 
myriandra Merr. 675 
nannodes Kosterm. 667 
nonda F.v.M. ex Bcnlh. 636, 

655, 658 map 
nonda auct. 658 
oblongifolia Hk.f. 636, 655, 

659, 660 map 
papuana C.T.Whilc 655, 658 

ssp. papuana 658, 659 

ssp. salomonense 
(C.T.White) Prance 
658, 659 map 

ssp. whilci Prance 658, 
659 map 
parva Koslcrm. 636, 655, 

657 map 
polyandra 64 1 
prancci Koslcrm. 656, 660, 

661 map 

rigida Kosierm. 656, 660, 

661 map 
rubiginosa Ridl. 663 
salomonense C.T. White 



Flora Malesiana 

[ser. I, vol. 104 


scabra Hassk. 641 
senegalensis DC. 641 
sumatrana (Jack) Benih. 

641, 656, 661 map, 

wallichiana R. Br. 665 
Parinari auct. 665, 675 
Parinarium Juss. 654 

sect. Cyclandrophora (Hassk.) 

C. Mucll. 665 
sect. Neocarya DC. 654 
sect. Petrocarya DC. 654 
subg. ///Hk.f. 665 
subg. Cyclandrophora 

(Hassk.) 31. 665 
subg. Macrocarya Miq. 

subg. Petrocarya (DC.) Miq. 

amboinense T. & B. 669 
asperulum Miq. 67 1 
borneense Merr. 659 
corymbosum (Bl.) Miq. 673 
costatum Bl. 

var. rubiginosum Ridl. 
curranii Merr. 669 
elaium King 669 
excelsum 654 
fragile T. & B. 646 
grifhihianum Benth. 673 
hahlii Warb. 669 
helferi Hk.f. 663 
heieropelalum Scort. ex King 

jackianum Benth. 670 
kunslleri King 677 
lanceolatum T. & B. 669 
laiifolium Hend. 666 
laurinum A. Gray 669 
maingayi King 671 
maranthes Bl. 673 
margarata A.Gray 669 
mindanaeruse Perk. 669 
mulliflorum (Korlh.) Miq. 

myriandrum Merr. 677 
nitidum Hk. f. 646 
nitidum auct. 674 
palauense Kaneh. 674 
philippinense Elmer 646 
polyneurum Miq. 664 
racemosum Vidal 674 
salicifolium (Presl) Miq. 674 
scabrum Hassk. 669 
spicaium King 671 
villamilii Merr. 671 
warburgii Perk. 669 

Parinarium auct. 661, 665 
Passiflora aurantia Forst. f. 717 
Passifloraceae 7: 405-434, 

829; 10: 166, 717 
Pedaliaceae 4: 216-221; 5: 

557; 7: 829 
/'e/ae Adanson 493 
Pentaphragmataceae 4: 517- 

Pentaphylacaceae 5: 121- 

124, 566 
Pentildis Zipp. ex Bl. 46 
Peponaster major Rumph. 94 
Pericampylus Miers 160, 169, 
170, 172, 227 
glaucus (Lamk) Merr. 165, 

228*, 229, 236 
incanus (Colebr.) Hk.f. & 

Th. 229 
lanuginosus (Bl.) Miq. 

membranaceus Miers 229 
Periomphale Baill. 335 
papuana Steen. 336 
Peripetasma polyanthum Ridl. 

Perrottetia alpestris (Bl.) Loes. 
ssp. philippinensis (Vidal) 
Ding Hou 716 
Petalinia Becc. 12 

bancana Becc. 14 
Petrocarya Schreb. 641, 654 
excelsa Jack 641, 670, 

glaberrima (Hassk.) Miers 

scabra (Hassk.) Miers 669 
sumatrana Jack 661 
Petrocarya auct. 665 
Petrosavia 109, 112 
Petrosaviaceae 109 
Peumus 259, 260-262 

boldus 260 
Phanerodiscus 4 
Phelima Nor. 605 
Pherosphaera 343 
Pherosphaeraceae 354 
Philbomea Hall.f. 607-609, 
magnifolia (SiapO Hall.f. 

614*, 615 map 
palawanica Hall.f. 615 
Philydraceae 4: 5-7; 7: 829 
Phlebocalymna 145 

lobospora F.v.M. 149 
Phyllocladaceae 354 
Phyllocladus L.C. Rich, ex 
Mirbel 337-343, 347, 
354, 355, 357 map 


hypophyllus Hk.f. 342. 
357*, 358*, 359*, 
var. protracta Warb. 359 
major Pilger 359 
protractus (Warb.) Pilger 359 
Phyllocosmus Klotzsch 621 
Phytocrene 91, 166 
/o/jm Merr. 219 
malacothrix Sleumer 227 
Phytolaccaceae 4: 228-232; 

5: 557 
Picea 338 
Pierotia Bl. 622 
lucida Bl. 625 
reticulata Bl. 626 
Pimela angustifolia Bl. 715 
Pinaceae 343, 347, 447-453 
Pinus L. 337-340, 342, 347, 
subg. Diploxylon 449 
dammara Lamb. 433, 434 
finlaysoniana Bl. 45 1 
insular is Endl. 452 
kasya Pari. 452 
kesiya Royle ex Gordon 341, 

346,451, 452 map 
khasia Engelmann 452 
var. insularis (Endl.) 
Gaussen 452 
khasyana Griff. 452 
latteri Mason 451, 452 
merkusiana Cooling & 

Gaussen 45 1 
merkusii Jungh. & de Vriese 
340, 341, 346,448*, 
449*, 450*. 451 map 
var. tonkinensis Chev. 
sumatrana Jungh. 451 
sylvestris auct. 45 1 
taeda auct. 452 
timorensis 452 
Piperaceae 127 
Piperales 127, 567 
Piptocalyx Oliv. ex Bth. 255, 
261, 326, 327, 330 
macrurus Gilg & Schltr 330, 

moorei Oliv. 330 
Pittosporaceae 5: 345-362; 
6: 960; 7: 829; 9: 568; 
10: 717 
Pittosporum moluccanum 

(Lamk) Miq. 717 
Platea 145 
Plumbaginaceae 4: 107-112 


Index to scientific plant names 


Podocarpaceae 338, 342, 343, 

347, 351-419, 420 
Podocarpus rHerit. ex Persoon 
337, 340, 343, 347, 351- 
355, 395, 397 map 
sect. Acuminatus de Laub. 

397, 398, 404 
sect. Dacrycarpus Endl. 374 
sect Dacrydioideae Bennett 

sect. Dacrydium Bertrand 374 
sect. Dammaroides Bennett 

sect. Foliolatus de Laub. 

397, 398, 399 

sect. Globulus de Laub. 397, 

398, 405 

sect Gracilis de Laub. 398, 

sect. Longifoliolatus de Laub. 

398, 407 
sect. Macrostachyus de Laub. 

398, 412 
secL Nageia Endl. 390 
sect. Polypodiopsis Bertrand 

sect. Polystachyus de Laub. 

398, 399, 416 
secL Prumnopiiys (Philippi) 

Bertrand 384 
sect. Rumphius de Laub. 

398, 399,406. 414 
sect Stachycarpus Endl. 

sect. Sundacarpus Buchholz 

& Gray 385 
sect. Taxoideae Bennett 384 
subg. Foliolatus de Laub. 

subg. Stachycarpus (Endl.) 

Engl. 384 
agalhifoliaB]. 391 
amara BI. 385 
archboldii N.E. Gray 399, 
402, 403 map 

var. crassiramosus N.E. 
Gray 409 
archboldii (non N.E. Gray) 

Gausscn 412 
atjchcnsis (Was.schcr) de 

Laub. 408 map. 409 
beccarii Pari. 393 
blumei EnA\. 391 
bomccnsis dc Laub. 398, 

400. 403 map 
bractcatus BI. 341,408,409 

var. brevipes BI. 408 


brassii Pilger 412, 413 map 

var. brassii 413 

var. humilis de Laub. 339, 
413, 414 
brevifolius (Stapf) Foxw. 

402,412,413 map. 414 
brevifolius (non StapO Foxw. 

celebica (non Hemsl.) Warb. 

celebicus Hemsl. 349 
cincius Pilger 383 
compacta Wasscher 383, 

confertus de Laub. 340, 341, 

407, 408, 409 map 
costalis PresI 412, 413 map, 

costalis (non Presl) Foxw. 

crassigemmis de Laub. 353, 

402, 412. 413 map 
cumingii Pari. 381 
cupressina sensu Lane-Poole 

cupressina R. Br. ex Mirbel 

cupressina Ridl. 379 

var. curvula Miq. 380 
dacrydiifolia Wasscher 383 
decipiens Gray 400 
deflcxusRidl. 340, 341.400, 

403 map 
discolor BI. 400 
dulcamara Seem. 387 
elaia R.Br. 453 
eurhyncha Miq. 385, 387 
f ale if or mis Pari. 372 
filicifolius Gray 395 
gibbsii N.E. Gray 398, 407, 

408 map 
glaucus Foxw. 409, 410. 

411 map 
globulus dc Laub. 406 map 
idenburgensis N.E. Gray 405 
imbricatus BI. 376 

var. cumingii (Pari.) 
Pilger 381 

var. curvula (Miq.) 
Wasscher 380 

var. kinabaluensis 

Wasscher 38 1 

imbricatus scnsu Foxw. 

imbricatus (non BI.) Gibbs 

379. 381 
insularis dc Laub. 399. 402. 

404 map 


javanica sensu Merr. 379 
javanicus (Burm. f.) Merr. 453 
javanicus (non Burm. f.) Merr. 

junghuhniana Miq. 400 
kawaii Hayata 379 
koordersii Pilger ex K. & V. 

latifolia f. ternaiensis de Boer 

latifolius (non Thunb.) BI. 

laubenfelsii Tiong 399. 415. 

416 map 
ledermannii Pilger 405 map 
leptophylla Wasscher 371 
leptostachya BI. 400 
levis de Laub. 398. 400, 

403, 404 map 
lophatus de Laub. 410, 411 

lucienii 406 
macrocarpus de Laub. 399, 

417, 418 map 
maximus (de Laub.) Gaussen 

micropedunculatus de Laub. 

339, 341. 405 map 
nakaii 406 
neglecta BI. 400 
neriifolius D.Don 399. 400 

var. atjehensis Wasscher 

var. bractcatus (BI.) 
Wasscher 408 

var. brevifolius Stapf 

var. brevipes (BI.) Pilger 

var. polyantha Wasscher 

var. ridleyi Wasscher 419 

var. teysmannii (Miq.) 
Wasscher 406 

var. timorcnsis Wasscher 
neriifolius D. Don in 

Lamb. 417 
neriifolius (non D. Don) 

Stecn. 408 
novaccalcdoniac 403 
palcmffanica Miq. 453 
papuanus Rki\. 379 
papuiinus (non Ridl.) Pilger 

papuanus (non Ridl.) Sicup. 



Flora Malesiana 

[ser. I, vol. 104 


pcdunculala Bailey 387 
philippinensis Foxw. 415 
piigcri Foxw. 402, 410. 411 

pilgeri (non Foxw.) v. Roycn 

polyanlha (Wasschcr) 

Gausscn 400 
polystachyus R.Br, ex Endl. 
339-341, 417*, 418 

var. rigidus Wasschcr 403 
polystachyus (non R.Br.) Li 

pscudobractealus dc Laub. 

342, 409 map 
ridleyi (Wasscher) N.E. Gray 

340, 341, 398,417,418 

map, 419 
ridleyi auct. 405 
rotundus de Laub. 410, 411 

rubcns de Laub. 399, 401*, 

402 map 
rumphii Bl. 403, 415,416 

schlechleri Pilger 410 
solomoniensis 403 
spathoides dc Laub. 398, 400, 

404 map 
steupii Wasscher 380, 381 

teysmannii Miq. 406 map 
ihevetiifolia B 1.417 
vitiensis Seem. 394 
wallichianus Presl 391 
wangii Chang 410 
Podocarpus Labill. 356 
Podostemaceae 4: 65-68; 

6: 963; 10: 717 
Polemoniaceae 4: 195-196 
Polyalihia pulchrinervia Boerl. 

Polycarpicae 165, 261 
Polygala L. 455-457, 459 
sect. Chamaebuxus DC. 455, 

460,461, 465, 467 
sect. Melchiora (Stccn.) 

Meijdcn 461, 465 
sect. Orihopolygala Chodat 

sect. Polygala 460, 461, 469 
sect. Pseudoscmciocardium 

Adema 460,461, 462 
sect. Semeiocardium [non 

(Zoll.) Hassk.] Chodat 

acicularis Oliv. 465 


arillala Buch.-Ham. 461, 

467*. 469 
arvensis Willd. 477, 478 

var. squarrosa Benth. 476 
arvensis (non Willd.) Adema 

arvensis (non Willd.) Benth. 

brachislachyos 476 
brachystachya BI. 477, 480 
brachystachya DC. 476, 

buchanani Buch.-Ham. ex 

D. Don 474 
buxiformis Hassk. 477 
cardiocarpa Kurz 461 , 463, 

cardiocarpa (non Kurz) Ridl. 

chamaebuxus 460 
chinensis (non L.) Benn. 477 
chinensis L. 458, 462, 477, 
478*, 479 

var. brachystachya (Bl.) 
Benn. 481 

var. linearifolia (non 
Willd.) Chodat 481 

var. triftora (L.) Benn. 482 
ciliata L. 487 
densiflora Bl. 478 
discolor Buch.-Ham. ex 

D. Don 474 
elongata WiUd. 477, 480 
elongata (non Willd.) Benn. 

criocephala Benth. 476 
eumekes Hassk. 481 
exsquarrosa Adema 460, 4o2, 

fernandesiana Paiva 475 
furcata Roylc 463, 465, 

glaucocarpa Ridl. 468 
glaucoides L. 462, 477, 479, 
480, 481* 

var. triflora (L.) Trimen 
glomerata Lour. 478, 479 
hondoensis Nakai 472 
humilis Span. 481 
isocarpa Chodat 463 
japonica Houtt. 462, 472* 
javana DC. 460, 461, 470*, 

karensium Kurz 469 
khasyana Hassk. 472 
leptalea DC. 474 
///iar//b//a Willd. 477, 481 


linarifolia (non Willd.) 

Adema 481 
linearifolia 48 1 
longifolia Poir. 460, 462, 

lutea 460 

luzoniensis Merr. 472 
macrostachya Hassk. 481 
malesiana Adema 461, 463, 

mariesii Hemsl. 465 
monspeliaca (non L.) Blco 

monticola HBK 460 
monticola (non HBK) Ridl. 

oligophylla DC. 474 
orcotrephes Burtt 461, 467* 
paenea 457 
palustris Lace 463 
paniculataL. 458, 460, 461, 

papuana (Steen.) Meijden 

461, 465, 466* 
persicariacfolia DC. 460, 

462, 473* 
polifolia Presl 462, 476, 

polyfolia 476 
polygama 458 
prostrata Willd. 482 
pulchra Hassk. 468 
pyramidalis Lev. 474 
rhinanthoides Benth. 460, 

461, 480* 
riukiuensis Ohwi 474 
rufa Span. 474 
septemnervia Merr. 474 
sibirica L. 473 
sibirica (non L.) Hassk. 472 
simadae Masam. 477 
simassan Miq. 468 
sumatrana Miq. 461, 467*, 

tatarinowii Regel 461, 462, 

464*, 488 
telephoides Willd. 477, 

telephoides (non Willd.) 

W. & A. 476 
tinctoria (non Vahl) Hassk. 

tonkinensis Chodat 469 
toxoptera Turcz. 478 
iranquebarica Mart. 478 
trichocolpa Chodat 469 
triflora L. 462, 479, 481, 



Index to scientific plant names 



lriner\aia Ham. ex Wall. 487 
iriphylla Buch.-Ham. ex 
D. Don 
var. glaucescens (non 
Wall.) Benn. 463 
triphylla (non Buch.-Ham. ex 

D. Don) Royle 462, 539 
umbonaia Craib 463 
undulaia Roxb. 487 
variabilis (non HBK) Hassk. 

vaulhieri Chodat 460 
venenosa Juss. ex Poir. 461, 
467*, 468 
ssp. pulchra (Hassk.) 

Sieen. 468 
ssp. venenosa Sleen. 469 
var. eramosa O.K. 468 
var. robusta Craib 468 
veronicaefolia All 
veronicea F.v.M. 472 
vulgaris L. 460 
vulgaris (non L.) Thunb. 472 
wallichiana Wight 474 
warburgii Chodat ex Warb. 

watiersii Hance 465 
wightiana W. & A. 460, 462, 
Polygalaceae 455-539, 639 
tribe Moulabeae 455, 458 
tribe Polygaleae 458 
tribe Xanihophylleae 458 
Polygalales 457, 458 
Polyosma 145 
Pontederiaceae 4: 255-261; 

5: 557; 6: 964 
Porotheca K.Sch. 187 

peliolaia K.Sch. 187 
Portulacaceae 7: 121-123 
Primulaceae 6: 173-192, 

Proteaceae 5: 147-206, 566; 
6: 965; 7: 830; 9: 568; 
10: 151 
Proteales 151 
Proicanae 151 
Proiium 123 

Protoalhcrospcrmoxylon 256 
Prumnopitys Philippi 343, 347, 
352, 353. 355. 384. 385 
sect. Sundacarpus (Buchhol/ 

& Gray) dc Laub. 385 
amara (Bl.) de Laub. 385. 
386*. 387 map. 388*. 

Prunus 640 

Pseudaleia imbricata (Roxb.) 
Hassk. ex Valeton 9 
ongistylis Hassk. ex 
Valeton 9 
Pleleocarpa Oliv. 29 
Pierocarya griffithiana (Benth.) 

Miers 674 
Pteroneurum DC. 550 
decurrens Bl. 554 
javanicum Bl. 551 
Ptychopeialum 2-4 
Punicaceae 4: 226-227 
Pycnarrhena Miers ex Hk.f. 
& Th. 157, 160-162, 
167-171. 172. 173, 178. 
auslraliana F.v.M. 175 
balabacensis Yamamoto 

balanensis Yamamoto 175 
borneensis Diels 1 75 
calocarpa (Kurz) Diels 174 
casianopsidifolia Yamamoto 

celebica (Boerl.) Diels 175 
elliptica Diels 175 
fasciculaia (Miers) Diels 

^ran^/5 K.Sch. & Uut. 175 
insignis (Hatus.) Forman 

173, 174 
longifolia (Dccne ex Miq.) 

Becc. 173, 174 
lucida (T. & B.) Miq. 173, 

174, 178 
manillensis Vidal 173, 175 
membranifolia Merr. 175 
merrillii Diels 175 
moniana Back. 174, 178 
nogovuineensis Miq. 173, 

ozantha Diels 173, 176*, 

177, 179 
papuana Kanch. & Hatus. 

plcniflora 173 
sayeri Diels 175 
lumcfacta Miers 159, 173, 

175, 176* 
Pyrolaccac 335 

Pyrrhosa horsfieldii (Bl.) Hassk. 

Pyrularia 33 

Qucrcus gilva Bl. 701 
var. procera Bl. 701 
jama-buwa Sicb. 699 

Radix puluronica Rumph. 

Ranunculales 163, 680 
RaphanusL. 541, 543,545, 
caudaius L. 546 
raphanistrum L. 547 
sativus L. 541, 546 
Reinwardlia 609 
Restionaceae 5: 416-420, 

Rhamnales 151 

Rhipogonum album R.Br. 253 
Rhizophoraceae 5: 429-493; 
6: 965; 8: 550; 9: 568; 
10: 253, 717 
Rhopalopilia 31-33 
pallens Pierre 33 
Rorippa Scop. 541, 543, 545, 
backeri (Schulz) Jonsell 541, 

555, 556 
benghalensis (DC.) Hara 555, 

dubia (Pers.) Hara 559 
heterophylla (Bl.) Williams 

555, 559 
hybosperma (Schulz) 

Jonsell 55, 559 
indica (L.) Hicm 

var. apeiala (Lour.) Hochr. 
islandica (Ocd.) Borb. 557 
micrantha (Roih) Jonsell 555, 

nasturtium-aquaticum (L.) 

Hayek 555, 556* 
officinalis (R.Br.) P. Royen 

palustris (L.) Becc. 555, 557 
pcekelii (Schulz) P. Roycn 

555, 557, 558* 
schlechteri (Schulz) P. Roycn 
Rosaceae 638, 639 

subfam. Chrysobalanoidcac 

subfam. Neuradoidcae 639 
subfam. Prunoidcae 640 
Rosalcs 639 
Roucheria 608 

coniesliana Pierre 619 
griffithiana Planch. 617 
Ruiaccac 165 kunstlcri King 539 

Sabah Mangtieiia Dandy ex 
Mcijcr 591 


Flora Malesiana 

[ser. I, vol. 104 

Sabia Colebrooke 163, 166, 
679-681, 682, 683 map 
campanulata Wall. 685 
celastrinea Muell. 685 
densiflora Miq. 690. 698 
elliplica (Miq.) Miq. 684 
crralica v. d. Water 683, 684 
floribunda Miq. 690, 698, 

harmandiana Pierre 687 
japonica 163 

javanica (Bl.) Backer ex Chen 
684, 685 
var. gtabriuscula (Bl.) 
Chen 684 
limonacea 685 

limoniacca Wall, ex Hk.f. & 
Th. 684, 685, 686*, 687* 
malabarica Bedd. 685 
menescorla 684 
menicosla 684 
meniscosia Bl. 684 

var. eltiptica Miq. 684 
\ai.firma Bl. 684 
var. glabriuscida Bl. 684 
var. latifoUa Bl. 684 
papuana Warb. 688 
parviflora Wall. 683, 687 
ssp. parviflora 685, 687 
ssp. philippinensis 
(Robins.) v.d. Water 
687, 688* 
var. harmandiana Lecomte 
pauciflora Bl. 683, 684, 688 
philippinensis Robins. 687, 

racemosa Chen 684, 689 
ssp. kinabaluensis 

v.d.Water 683, 689 
ssp. racemosa 689 
reticulata Elmer 688 
sumatrana Bl. 682, 683, 689 
Sabiaceae 163, 166,679-715 
Salacia banlettii RidJ. 25 
Salicaceae 5: 107-110 
Salmonea 486 
Salomonia Lour. 455-457, 
459. 486 
sect. Epirixanthes (Bl.) Benn. 

angulata Griff. 488 
aphylla Griff. 490 
arnottiana Miq. 488 
canarana Miq. 488 
cantoniensis Lour. 487 
cantoniensis auct. 488 
cavalereriei Lev. 488 
ciliata (L.) DC. 487 


cordata Wight 487 
edentula DC. 487 
elongata (Bl.) Kurz ex Koord. 

horneri Hassk. 488 
longiciliata Kurz 488 
mariinii L^v. 488 
oblongifolia DC. 487 
obovata Wight 487 
parasitica Griff. 490 
peiiolata D. Don 487 
ramosissima Turcz. 488 
rigida Hassk. 488 
seguinii Lev. 488 
sessiliflora 487 
sessilifolia D.Don 487 
setoso-ciliata Hassk. 488 
stricta Sieb. & Zucc. 488 
subrotunda Hassk. 487 
trinervata Steud. 487 
uncinata Hassk. 488 
Salomonia auct. 488 
Salvadoraceae 4: 224-225 
Sampacca O.K. 598 

domestica IV alba Rumph. 

longifolia O.K. 605 
monlana O.K. 603 
montana Rumph. 582 
sylvestris Rumph. 596 
velutina O.K. 603 
Santalaceae 2, 3, 33-35, 52 
Santalales 5. 34 
Sapindaceae 639, 679, 680, 681 
Sapindales 163,458.639,681 
Sapindus microcarpus W. & A. 

Sapotaceae453, 593 
Sarcandra Gardner 123. 125. 

126. 128, 129, 134, 144 
glabra (Thunb.) Nakai 124, 
127, 128, 134, 135*, 
ssp. brachystachys (Bl.) 
Verdcourt 136 
var. brachystachys 136 
var. melanocarpa (Ridl.) 
Verdcourt 136 
hainanensis (Pei) Swamy & 

Bailey 136 
glabra auctt. mult. 136 
Sarcodiscus Griff. 287 

chloranthiformis Griff. 298 
Sarcopetalum F.v.M. 157-161, 
167, 169. 172, 224 
harveyanum F.v.M. 159, 
224. 228* 

Sarcosperma 29 
Sarcosperma{ta)ceae 4: 32- 

34; 6: 967; 10: 29 
Sarcostegia Benih. 641 
Sanima53. 57, 59,61-64 

henryi 62 
Saururaceae 4: 47—48 
Saxegothacaceae 354 
Saxifragaceae 145 
Scaphocalyx parviflora Ridl. 

spathacea Ridl. 716 
Schisandraceae 163, 679 
Schoepfia Schreb. 1-6, 27 

fragrans Wall. 2, 27, 28* 
Schoepfiopsis Micrs 27 

fragrans (Wall, in Roxb.) 
Miers 27 
Sciadocarpus Hassk. 287 

brongniartii Hassk. 298 
Sciadopitys 354 
Sciaphila Bl. 110, 111 map, 

112, 720 

sect. Hermaphroditantha 
subsect. Polyandra 112 

sect. Hexanihera 1 13 

sect. Oliganthera 

subsect. Bilobatae 113 
subsect. Quadrilobatae 

112, 113 
subsect Trilobatae 

affinis 'Qtcc. 113 

andajensis Becc. 117 

arfakiana Becc. 109, 112, 

113, 115*. 117 
aster ias Ridl. 116 
atroviolacea Schltr 118 
australasicaWQmsX. 117 
brachystyla Schltr 1 16 
buruensis 112 
clemensae Hemsl. 117 
conferta J.J. Smith 115 
consimilis Bl. 113, 120 
corallophyton K.Sch. & 

Schltr 113, 119 

var. gracilis Giesen 1 19 
comiculata Becc. 112, 115, 

116, 117 
corniculata (non Becc.) Went 

crinita Becc. 117 
decipiens Backer 113 
densiflora Schltr 112, 113, 

118, 119* 
dolichostyla Schlti 119 
erubescens (Champ.) Miers 



Index to scientific plant names 



flexuosa Giesen 118 
galiensis Schllr 115 
hermaphrodila Schltr 1 13 
hydrophila Schllr 120 
inaequalis Schltr 1 18 
inornaia Fetch 116 
long i pes Schltr 118 
maboroensis Schltr 1 16 
macra K.Sch.&Uut. 120 
macro K.Sch. & Schltr 

macra Schltr 120 
maculata Micrs 112, 113 
major Bccc. 116 
micranthera Giesen 112, 121 
mindanaensis G'xcscn 120 
minuta Schltr 113 
moniicola K.Sch. & Schltr 

multiflora Giesen 112, 113, 

nana BI. 113, 116, 117 
neo-caledonica Schltr 1 15 
nu/an.y Giesen 118 
oligochaeie Schltr 1 15 
papillosa Becc. 121 
papuana Becc. 116 
pilulifera Schllr 1 16 
pumila Giesen 1 14 
purpurea 109 
quadribuUifera J.J.Sm. 112, 

reficxa Schlu 118 
sccundiflora Thw. ex Bih. 

112, 115*. 116, 120 
slcmmermannii Fosb. & 

Sachet 120 
suhhermaphrodita J.J. Sm. 

sumairana Becc. 1 16 
tcnclla Bl. 112. 113, 115* 

var. robusta Giesen 1 14 

var. voigiii Giesen 1 14 
torricellensis K.Sch. & Schllr 

trichopoda Schltr 1 1 8 
tuber culata Giesen 112, 118 
valida Giesen 1 18 
versteegiana Went 1 16 
viliensis A.C. Smith 118 
wariana (Schltr) 

Mccrcndonk 112, 117 
werneri Schltr 1 16 
wmklcri Schlu 112. 113. 

Sclcropyrum 33 

aurantiacum (Laut.& K.Sch.) 

Pilgcr 52 


pentandrum (Dennst.) 

Mabberley 52 
wallichianum (W. & A.) Am. 
Scorodocarpus Becc. 1-6, 14 
map, 15 
bomeensis (Baill.) Becc. 5. 
15, 16* 
Scyphosiegia Stapf 326 
Scyphostegiaceae 5: 297- 299; 
6: 967; 7: 830; 10: 326 
Sccuridaca L. 455-459, 483 
atro-violacea Elmer 483, 484 
bracieata Benn. 484 

var. papuana F.v.M. 485 
complicata auct. 484 
corymbosa Turcz. 484 
cumingii Hassk. 484 
ecristata Kassau 483, 485* 

var. niiida Kassau 485 
inappendiculata Hassk. 483 
ssp. corymbosa (Turcz.) 

Meijden 484 
ssp. inappendiculata 484 
paniculaia Roxb. 484 
philippincnsis Chodat 483, 

scandens Ham. ex Benih. 

lavoyana Wall, ex Benn. 484 
volubilis auct. 484 
yaoshannensis Hao 484 
Selwynia F.v.M. 218 
laurina F.v.M. 219 
Semeiocardium Zoll. 539 
hamillonii Hassk. 463 
Semeiocardium (non Zoll.) 

Hassk. 459, 462 
Shepherdia Null. 151 
Simaroubaceae 6: 193-226, 

968; 10: 621, 622, 718 
Sinapis 541 
alba L. 545 
juncea L. 546 
imoriana DC. 546 
Siparuna 259 
gilgiana 261 
guyanensis 261 
Siparunaccae 261 
Siph()n(Kl()n celasuincus Grill. 

Sisymbrium 541 , 543 
amphibujm L. 

var. pcduslre L. 557 
micranihum Rolh 560 
naslurtium-aqualirum L. 555 
Skaphium Miq. 493 
anceatuin Miq. 527 

Sonneratiaceae 4: 280-289, 

513; 5: 557; 6: 973 
Sparganiaceae 4: 233-234; 

10: 718 
Sparganium fallax Graebn. 718 
simplex Huds. 

f. simplex 718 
subglobosum Morong 718 
Spermabolus T. & B. 605 

fruticosa T. & B. 605 
Sphaerocarya leprosa Dalz. 22 
Sphenocleaceae 4: 27-28 
Sphenostemon Baill. 145, 146, 
326, 333 
sect. Apetalae (Stecn.) 

Steen. 147 
sect. Sphenostemon 147 
scr. Apetalae Stecn. 147 
ser. Sphenostemon 146, 147 
arfakensis (Gibbs) Steen. & 

Erdtman 147, 149 
lobosporus (F.v.M.) L.S. 

Smith 147, 149 
oppositifolius Hiirl. 146 
pachycladus 146 
papuanus (Laul.) Stecn. & 

Erdtman 147, 148*, 149 
pauciflorum (A.C. Smith) 
Steen. & Erdtman 147, 
Sphenostemonaceae 145-149, 

326, 333 
Siachycarpus (Endl.) Ticgh. 352, 
sect. Sundacarpus (Buchholz 

& Gray) Gausscn 385 
amara Gausscn 387 
Stackhousiaceae 4: 35-36 
Staphyleaceae 6: 49-59 
Stcganthcra Perkins 255-263, 
288, 306, 321 
alpina Perkins 319 
atepala Perkins 319 
austral iana C.T.White 309. 

brassii (A.C. Smith) Kaneh. 

& Hatus. 315 
buergersiana Perkins 316 
chimbucnsis Philip.son 309, 

crispula Perkins 316 
cyclopcnsis Philipson 308. 

denlaliJ (Val.) Kanch. & 

Hatus. 308, 310 
clliptica A.C. Smith 278 
fasciculata Philipson 307*. 

308. 309 
fengcriana F'crkins 315 


Flora Malesiana 

[ser. I, vol. 104 


forbesii Perkins 316 
hcntyi Philipson 308, 309 
hirsuta (Warb.) Perkins 262, 

300, 308.309, 314*, 

315, 318 
hospilans (Becc.) Kaneh. & 

Hatus. 257, 258, 308- 

310 311, 313*. 319 
ilicifolia A.C. Smith 256, 

296, 309, 317*, 318 
insculpta Perkins 308, 309, 

insignis Perkins 312 
ledcrmannii (Perkins) Kaneh. 

& Hatus. 257, 308, 319 
moszkowskii (Perkins) 

Kaneh. & Hatus. 257, 

308, 309, 311 
myrtifolia (A.C. Smith) 

Philipson 309, 313 
oblongiflora Perkins 315, 

oblongifolia 315 
odontophylla Perkins 319 
oligantha (Perkins) Kaneh. & 

Hatus. 308, 309, 315 
oligocarpella Kaneh. & 

Hatus. 291 
parvifolia (Perkins) Kaneh. & 

Hatus. 309, 315 
psychotrioides Perkins 319 
pycnoneura Perkins 319 
riparia Kaneh. & Hatus. 316 
royenii Philipson 257, 308, 

309, 310 
salomonensis (Hemsl.) 

Philipson 308, 309, 311, 

schlechleri Perkins 278 
schumanniana Perkins 3 1 5 
suberoso-alata Kosterm. 311 
symplocoides Perkins 319 
thyrsiflora Perkins 315, 316 
lorrlcellensis Perkins 316 
villosa Kaneh. & Hatus. 

warburgii Perkins 315 
Stcllatopollisbarghoomii 256 
Sicmorwaws fruiescens Bl. 25 

membranaceus Bl. 22 
Stcphania Lour. 157, 160-162, 

165, 167, 169, 171, 243 
sect. Eustephania Diels 243, 

sect. Thamnoihyrsa Diels 

243, 244 
acuminaiissima (Bl.) Spreng. 



borneensis Yamamoto 

capitata (Bl.) Spreng. 159, 

160, 161, 165, 167,236, 

243-245, 252*, 253 
calosepala Diels 249, 250 
calosepala (non Diels) Merr. 

cauliflora Becc. 249, 250 
cincinnans K. Sch. 203 
concinna Miers 247 
corymbosa (Bl.) Walp. 243- 

245, 249, 250 
corymbosa (non Bl.) Turcz. 

dictyoneura Diels 160, 167, 

243, 244, 253 
discolor (Bl.) Spreng. 245 

var. hernandiifolia (Willd.) 
Boerl. 245 
exigua Miers 247 
florulenta Becc. 251 
formanii Kundu & Guha 250, 

forsteri (DC.) A. Gray 247 
glaucescens (Decne) Walp. 

grandiflora Forman 244, 248 
hallieri Diels 247 
hernandiifolia (Willd.) Walp. 
245, 247 

var. discolor (Bl.) Miq. 

var. genuina (Bl.) Miq. 

var. glabra (Bl.) Miq. 245 
hernandiifolia [non (Willd.) 

Walp.] Koord. 249 
hernandiifolia [non (Willd.) 

Walp.] Ridl. 248 
hernandiifolia [non (Willd.) 

Walp.] Vidal 251 
japonica (Thunb.) Miers 159, 
165, 243-245, 245, 
246*, 247 

var. discolor (Bl.) Forman 
247, 248 

var. japonica 247 

var. timoriensis (DC.) 
Forman 247 
japonica [non (Thunb.) 

Miers] Miers 25 1 
longifolia Becc. 252 
mcnadonensis Diels ex 

Koord.-Schum. 249 
merrillii Diels 249, 250 
moluccana Forman 244, 245, 



montana Diels 159, 243- 

245, 250, 253 
ncoguineensis Kundu & Guha 

obvia Miers 252 
prapaiensis Yamamoto 248 
psilophylla (Presl) Forman 

243-245, 251 
ramosii Diels 249 
ramuliflora Miers 249 
reticulata Forman 244, 245, 

rotunda (non Lour.) Miq. 

rotundifolia 248 
salomonum Diels 244, 245, 

tomentosa (Bl.) Spreng. 

truncata Yamamoto 252 
venenosa 248 

venosa (Bl.) Spreng. 243- 
245, 248 
zippeliana Miq. 243-245, 

Strakaea Presl 65 

melastomaefolia Presl 78 
Strombosia Bl. 1-6, 19 

ceylanica Gardn. 20*, 21, 22, 
23 map 

var. lucida (T. & B. ex 
Valet.) Hochr. 22 

var. membranacea (Bl.) 
Hochr. 22 

var. sessilis Hochr. 22 
dubia Vidal 22 
elmeri Salvosa 22 
javanica Bl. 5, 20*, 21 map 

var. sumatrana Val. 21 
javanica auct. 22 
lucida T. & B. 22 
maingayi (Mast.) Whitm. 22 
membranacea (Bl.) Valet. 22 
minor Elmer ex Merr. 22 
mulliflora King 22 
philippinensis (Baill.) Rolfe 

21, 22 
philippinensis [non (Baill.) 

Rolfe] Lam & Holthuis 29 
rapaneoides S. Moore 22 
rotundifolia King 22 
Strombosiopsis 3, 4 
Slropha Noronha 129 
Stylidiaceae 4: 529-532; 

5: 564; 6: 976 
Styracaceae 4: 49-56; 6: 

976; 9: 568 


Index to scientific plant names 


Symplocaceae 8: 205-274; 
9: 569; 10: 718 

Symplocos Jacq. 

subg. Hopea Clarke 718 

ambangensis Noot. 719 

cochinchinensis 719 
ssp. laurina 719 

cosialifrucia Noot. 719 

hcrzogii 718 

iliaspaiensis Noot. 719 

laeteviridis 719 

maliliensis 719 


ssp. cumingiana 719 

rayaeNoot. 718, 719 

riangensis Noot. 718, 719 

vcrtJciliifolia 718 
Synandrium exsertum 243 

inclusum 243 

Taccaceae 7: 806-819 
Talauma Juss. 563, 568 
sect. Aromadendron Nliq. 

569, 576 
angatensis (Blco) Vidal 585 
aihlianiha Dandy 583 
beccarii Ridl. 586 
betongensis Craib 585 
binluluensis Agostini 577 
horneensis Merr. 583 
candollei auct. 585 
candollii Bl. 569,581,582 

var. laiifolia Bl. 582 
elegans (Bl.) Miq. 577 

var. glauca (Korth.) 
Parmcnt. 577 
elegans auct. 583 
elmeri Merr. ex Soderbcrg 

forbesii King 582 
giganiifolia Miq. 586 
giganiifolia auct. 585 
gilingensis Elmer 583 

var. glabra Dandy 583 

var. roiundaia Dandy 
glaucum (Korth.) Miq. 577 
gracilior Dandy 583 
grandiflora Merr. 585 
injlata Parment. 582 
intonsa Dandy 588 
javanica Parment. 583 
kunstlen King 582 
kuleinensis Agostini 586 
lanigera auct. 586 
levissinui Dandy 585 
Uliifcra {L.)O.Y.. 582 


longifolia (Bl.) Ridl. 582 
luzoniensis Warb. ex Perkins 

macrophylla Bl. 583 
magna Agostini 586 
megalophylla Merr. 586 
miqueliana Dandy 582 
mutabilis Bl. 582 

var. acuminata Bl. 582 

var. acuminaiissima 
T. & B. 583 

var. brevifolia T. & B. 

var. laiifolia T. & B. 583 

var. leiocarpa T. & B. 

var. longifolia Bl. 582 

var. sciagraphia 582 

var. splendens Bl. 582 
mutabilis auct. 585 
oblanceolata Ridl. 585, 586 
oblongata Merr. 585 
obovaia Korth. 585 
obovata auct. 586 
oreadum Diels 583 
avails Miq. 596 
papuana Schltr 596 
peninsular is Dandy 583 
persuaveolens Dandy 587 
plumierii (Schwartz) A. DC. 

pubescens Merr. 596 
pumita Bl. 582 
pumila auct. 583 
rabiana Craib 

var. villosa (Miq.) 
Parment. 588 
reticulata Merr. 583 
rubra Miq. 582 
rumphii Bl. 582 
sarawakensis Agostini 588 
sclerophylla Dandy 585 
sebassa Miq. ex Dandy 582 
singapurensis Ridl. 586 
soembensis Dandy 583 
sumulrana Agostini 583 
undulatifotia Agostini 583 
villariana Rolfc 583, 585 
villarii 583 
villosa Miq. 588 

f. celebica Miq. 603 
vrieseana Miq. 569, 596 
Tambouri.ssa 257, 258 

ficus(Tul.) A.DC. 326 
Taxaceae 343, 346, 347-35 1 . 

Taxaics .347 
laxodiaccac 338, 343, 354 

Taxus L. 337, 342, 347, 348 
baccata (non L.) Mast. 349 
ssp. cuspidata 

var. chinensis Pilger 
ssp. wallichiana (non 

Zucc.) Pilger 349 
var. sinensis Henry 349 
celebica (Wall.) Li 350 
chinensis (Pilger) Rehd. 350 
cuspidata (non Sieb.& Zucc.) 

Kaneh. 350 
cuspidata var. chinensis (Pil- 
ger) Rehd. & Wilson 350 
mairei (Lemee & Lev.) Hu & 

Liu 350 
speciosa Florin 350 
sumatrana (Miq.) de Laub. 

349, 350*, 351 map 
wallichiana (non Zucc.) 

Foxw. 349 
wallichiana var. chinensis 

(Pilger) Horin 350 
yunnanensis Cheng 350 
Temstroemiaceae 327 
Tetanosia Rich, ex M. Roemer 
olacioides (W. & A.) 
M. Roemer 47 
Tctrastylidium 3,4 
Tetrasynandra 284 map 
Thalamia Spr. 356 
Thalictrum 165 
Thcaceae 145 
Thealcs 145 
Thottea Rottb. 53-55, 57-59, 
61-64, 65 
beccarii Ding Hou 67, 74 
bomeensis Val. 65, 66, 73 
celebica Ding Hou 67, 78, 

corymbosa (Griff.) Ding Hou 
56*, 57, 59, 62, 67, 78, 
curvisemcn Ding Hou 61, 66, 

depcndens (Planch.) Klotzsch 

56*, 57, 62, 67, 75 
dinghoui 62 
grandillora Rottb. 59, 62. 66, 

68*, 69 
macTdnlha (Boerl.) Ding Hou 

60*, 62. 65, 67. 81 
macrophylla Bccc. 5'), 66, 71 
muluensis Ding Hou 56*, 
57, 67. 76*, 78 


Flora Malesiana 

[ser. I, vol. 104 


parviflora Ridl. 59, 62, 66, 

67, 80* 
paucifida Ding Hou 59, 62, 

67, 72*, 73, 81 
pcniiilobala Ding Hou 57, 

67. 76*, 77 
philippincnsis Quis. 67, 75, 

rcniloba Ding Hou 60*. 67, 

rhizantha Bccc. 67, 75 
robusta Siccn. 66, 70*, 71 
siliquosa (Lamk) Ding Hou 

sp. Ding Hou 83 
sU'aalmanii Ding Hou 66, 69 
sumau-ana (Mcrr.) Ding Hou 

67, 80*. 81 
lomcntosa (Bl.) Ding Hou 

54, 57, 59, 62, 65, 79 
iricornis Maingay 62. 67, 

74, 80* 
iriscrialis Ding Hou 55, 59, 

66, 69 
Thuja 444 

javanica Burm.f. 453 
papuana (F.v.M.) Voss 444 
Thujoidcae 444 
Thuya javanica 453 
Thymelaeaceae 4: 349-365; 

6: 1-48,976; 7: 830; 

10: 151 
Thymclacalcs 151 
Tiliacora Colcbr. 157, 160-168. 

170, 172, 185 
triandra (Colcbr.) Dicls 162, 

166, 185. 187, 208* 
Tinomiscium Micrs 157-162, 

166-170. 204 
arfakianum (Bccc.) Dicls 

197 • 
elasticwn Bccc. 205 
javanicum Micrs 205 
molle Dicls 205 
pctiolare Hk.f. & Th. 159, 

205. 206* 
phlUppinense Dicls 205 
phytocrenoides Kur/. ex 

T. & B. 205 
pyrrhoboiryum Miq. 205 
Tinospora Micrs 19, 158-171, 

andamanica Dicls 199 
angusta 190 
arfakiana Bccc. 190, 191, 

bacnzigeri Forman 159, 160, 

190. 191. 195, 196*, 197 


cclcbica Dicls 190, 193, 194 
cordifolia 190, 197 
coriacea (Bl.) Bcunndcex 

Hcync 199 
crispa(L.)Hk.f. & Th. 160, 

165, 166, 190, 191, 194, 

195, 196*, 197, 200 
cmpa [non(L.) Hk.f. &Th.] 

Dicls 199 
curtisii Ridl. 253 
dcniata 190 
dissitiflora (Laut. & K.Sch.) 

Dicls 159, 190, 191. 

192*. 197 
glabra (Burm.f.) Mcrr. 159, 

165, 186*. 191. 192*, 

194, 195. 199. 200 
glandulosa Mcrr. 190. 194 
hasiala Elmer 193 
havilandii Dicls 193,210 
hirsuia (Bccc.) Forman 190, 

homoscpala Dicls 190, 200 
macrocarpa Dicls 191, 192*, 

198, 199 
megalobotrys Laut. & 

K.Sch. 187 
mcrrilliana Dicls 159, 190, 

191, 191 
minuliflora K. Sch. & Laut. 

negrotica Dicls 1 93 

var. moniicola Yamamoto 
peekelii Dicls 197 
polygonoides Dicls 200 
pseudo-crispa Bocrl. 199 
reticulata 199 
rumphii Bocrl. 194. 195. 

sagitlata 190 
sinensis (Lour.) Mcrr. 190, 

smilacina Benth. 191, 196* 
subcordata (Miq.) Dicls 159. 

191, 192*, 200 
sumau-ana (Schcff.) Bccc. 
190, 191, 198, 199 

var. hanadac Yamamoto 
icijsmannii Bocrl. 191, 192*. 

tinosporoidcs (F.v.M.) 

Forman 190, 192* 
trilobata Dicls 159, 190, 

191, 202 
tuberculata (Lamk) Beumee 

ex Hcync 194 


uliginosa Micrs 199 
uliginosa (non Micrs) Hk.f. 
& Th. 198 
Trapaceae 4: 43-44; 6: 982 
Trcmandraccac 458 
Triccrcandra A. Gray 128 
Trichocarya Miq. 645, 648 

splendens 645 
Triclisia 165 
Trigoniaceae 4: 58-60; 10: 

Trimcnia Sccmann 145, 255, 
261, 326. 327, 329 map 
arfakensis Gibbs 330, 331 
bougainvilleensis (Rodcn- 
burg) A.C. Smith 332 
grandifolia Warb. 333 
macrura (Gilg & Schllr) 
Philipson 329 map, 330, 
331, 332 
moorei (Oliv. in Bth.) 
Philipson 328, 329 map, 
myricoides Gilg & Schltr 

330, 331 
ncocalcdonica 328, 329 map. 

papuana Ridl. 328*, 329 

map, 330, 331* 
weinmanniifolia Seem. 328, 
329 map, 330-332 
ssp. bougainvilleensis 
Rodenb. 328, 330, 332 
Trimeniaceae 145. 255. 261. 

Trimeria grandifolia (Hochst.) 

Warb. 333 
Trimeriza Lindl. 65 
Trisiichocalyx F.v.M. 217 
Triuridaceae 109-121. 110 

map, 720 
Triuridalcs 109 
Triuris 110, 112 
Trochiscus Gilli 555 

macrocarpus Gilli 559 
Tropacolaccac 639 
Tsjeru-caniram Rhecde 49 
Tsjerucaniram 49 
Tsjerucanirum 49 
Tsoongiodendron 563. 564. 567, 

Tsuga 338, 343 

mairei Lcmee & L6v. 350 
Tuba baccifera Rumph. 213 

flava Rumph. 210 
Turneraceae 4: 235—238 
Typhaceae 4: 242-244; 6: 


Index 10 scientific plant names 


Ulmaceae 8: 31-76, 551; 

9: 569; 10: 29 
Umbelliferae 4: 113-140, 

595; 5; 555; 6: 983: 

7: 830: 9: 569 
Uroboirya Stapf 31-35, 41 
sccL Lepionuroides Hiepko 

42 map 
sect. Urobotr>'a 42 
floresensis Hiepko 42 map, 

laiisquama (Gagn.) Hiepko 

32, 43, 42 map 
longipes (Gagn.) Hiepko 

42 map 
parv'iflora Hiepko 41*, 42 

map, 43 
siamensis Hiepko 34, 42 

map, 43, 46 
Uvaria-leaved Magnolia Meijer 


Vaccinium angulaium J.J. Sm. 

commuialum Mabberley & 

Sleumer 716 
Valerianaceae 4; 253-254 
Vallisneria 112 
VanhalUa Schuli. 65 
omenlosa J. A. &. J.H. 

Schultes 79 
V'idara liuorea Rumph. 1 1 
Viola rheophila Okamoio 720 
Violaceae 7: 179-212,831; 

10: 720 
Violaria Post & O.K. 568, 569 
Viscaceae 353 
Vochysiaceae 458 

Wcinmannia 123 
Weltingionia arnottiana Mcisn. 

Wilkiea F.v.M. 255-257, 262, 
263. 282, 288 
forcmanii Philipson 282, 

hucgciiana 288 
macrophylia 288 
WitLslcmia F.v.M. 335 

papuana (Stccn.) Siccn. 336* 
Worcesierianihus Men. 29 

Xanihophyllum Roxb. 455-459. 
subg. Brunophyllum Mcijdcn 
494. 535 


subg. Coriaceum Meijden 

494, 528 
subg. Exserium Meijden 494, 

subg. Triadelphum Meijden 

494, 530 
subg. Xanthophyllum 494, 

secL Eystathes (Lour.) 
Meijden 500, 507 
subsect. Eystathes 507, 

subsect. Jakkia (Bl.) 
Meijden 507, 508 

sect. Xanthophyllum 500 
acuminaiissimum Miq. 515 
adenopodum Miq. 501 
adenoius Miq. 497, 498, 511, 

var. adenotus 516 

var. Imeare Meijden 516 
affine Korth. ex Miq. 498, 
500, 501, 503 

B excelsa K. & V. 500 

a genuina K. & V. 503 

var. adenopodum (Miq.) 
K. & V. 501 
affine (non Miq.) K.Sch. &; 

Hollr. 537 
affine (non Miq.) Koord. 5 1 2 
affine (non Miq.) Ridl. 528 
amoenum Chodat 499, 533 
amoenum (non Chodat) Keith 

ancolanum Miq. 498, 511 

f. angustifolia Miq. 5 1 1 
angustigcmma Meijden 496, 

arsaiii C.E.C. Fischer 516 
bcccarianum Chodat 495, 

bomeense Miq. 495. 508 
bractealum Chodat 518*. 

496, 519 
brevipcs Meijden 495. 535, 

brigittae Mcijdcn 496, 498, 

511, 512* 
bullatum King 498. 501 
cclcbicum Meijdcn 499, 532 
ccraccifohum Mcijdcn 497. 

chartaccum Mcijdcn 499, 

535, 539 
ciirifoltum Chodat 530 
clovis (Slccn. ex Mcijdcn) 

Mcijdcn 496. 517 
cockbumii Mcijdcn 494, 502 


contracuim Meijden 499, 

cordaium Korth. ex Miq. 515 

f . aequale Chodat 5 1 5 
curtisii King 514 
densiflorum Chodat 521, 522 
discolor Chodat 495, 520 
ssp. discolor 520 
ssp. macranihum Meijden 
ecarinatum Chodat 499, 539 
ellipticum Korth. ex Miq. 

499, 530, 531* 
er>'lhrosiachyum Gagnep. 

497, 511, 523 
eiu-hynchum Miq. 497, 524. 
ssp. eurhynchum 525 
ssp. maingayi (Hk. f. ex 
A.W. Benn.) Meijden 
excelsum Bl. 503 

var. affine (Miq.) Boerl. 
excelsum (Bl.) Miq. 500 
ferrugineum Meijden 498, 

flavescens (non Roxb.) 

F.-Vill. 514 
flavescens Roxb. 498, 500, 
5(W, 528 
var. virens (Roxb.) 
A.W. Benn. 528 
flavovirens Elmer 520 
flavum Ridl. 505, 507 
floriferum Elmer 501 
forbesii Baker 523 
glabrescens Ridl. 508 
glandulosum Merr. 501 
glaucescens Miq. 539 
glaucum Wall, ex Hassk. 527 
gracile Chodat 5 1 3 
griffithii Hk.f.ex A.W. Benn. 
496, 513, 514 
ssp. angustifolium (Ng) 

Meijdcn 513 
ssp. crecium Meijden 513. 

var. angustifolium Ng 5 1 3 
var. curiisu (King) Ng 

var. montanum Ng 5 1 3, 
griffiihii (non A.W. Benn.) 

hchecarpum Chodat 539 
hctcrophyllum Mcijdcn 496, 


Flora Malesiana 

[ser. I, vol. 104 


heteropleurum Chodat 505, 

hildebrandii Meijden 498, 

hookerianum King 514 
hosei Ridl. 498, 502 
hypoleucum Merr. 520 
impressum Meijden 496, 

inccrium (Bl.) Meijden 496, 

insigne A.W. Benn. 536 
kalimantanwn Meijden 539 
kingii Chodat 530 
korthalsianum Miq. 496, 

kunslleri King 514 
laeve Meijden 499, 524 
lanceatum (Miq.) J.J. Sm. 

494, 496, 527 
lanceolatum 527 
latcriflorum Miq. 497, 527 
loheri Merr. 501 
longifolium (Bl.) Dietr. 514 
macranlhum Chodat ex Elmer 

macrophyllum Baker 498, 

maingayi Hk. f . ex 

A.W. Benn. 524, 525 
malayanum Meijden 495, 

microcarpum Chodat 527 
molle Ridl. 522 
montanum Meijden 499, 532 
moniicolum Meijden 496, 

muliiramosum Elmer 501 
ncglectum Meijden 497, 498, 

ngii Meijden 496, 526 
nigricans Meijden 495, 508 
novoguineense Meijden 496, 

obscurum A.W. Benn. 499, 

ovatifolium Chodat 498, 508 
palawancnse Elmer 497, 516 
palembanicum Miq. 5 1 1 
palembanicum (non Miq.) 

Keith 509 
palembanicum (non Miq.) 

King 524. 525 

pallidum Ridl. 501 
paniculatum Miq. 514 
papuanum Whitm. ex 

Meijden 457, 499, 537, 

parvifolium Meijden 496, 

parvum Chodat 5 1 3 
pauciflorum Meijden 496, 

pedicel latum Meijden 495, 

penibukanense Heine 495, 

petiolatum Meijden 496, 517 
philippinense Chodat 498, 

pseudoadenotus Meijden 495, 

pseudostipulaceum Merr. 5 1 3 
pseudoslipulaceum (non 

Merr.) Meijer 519 
puberulum Ridl. 525 
pubescens Meijden 495, 

pulchrum King 495, 521, 

ssp. pulchrum 521 

ssp. siapfii (Chodat) 
Meijden 522 
purpureum Ridl. 495, 522 
ramiflorum Meijden 494, 

495, 529*, 530 
reflexum Meijden 496, 519 
resupinatum Meijden 498, 

reticulatum Chodat 495, 

retinerve Meijden 497, 524 
robustum Chodat 515 

var. elmeri Chodat 5 1 5 
rufum Benn. 494, 505, 

sarawakense Chodat 503 
saraH'akensis 503 
schizocarpon Chodat 495, 

scortechinii King 536 
spec. 504 

spec. Anderson 530 
spec. £■ Ng 539 
stapferi 521 
5/apyz/ Chodat 521, 522 


stipitatum A.W. Benn. 497, 
499, 535 

var. borneense Chodat 535 

var. glabrum Meijden 535 

var. nitidum Chodat 533 

var. pachyphyllum Chodat 

var. stipitatum 535 
subcoriaceum (Chodat) 

Meijden 498, 509 
suberosum C.T.White 499, 

533, 534* 
subglobosum Elmer 539 

var. longifolium Elmer 
sulfureum 502 
sulphureum King 494, 502 
sumatranum Miq. 51 1 
tardicrescens Meijden 498, 

tenue Chodat 498, 508 
tenuipetalum Meijden 498, 

trichocladum Chodat 495, 

velutinum Chodat 495, 505 
venosum King 497, 525 
verrucosum Chodat 524, 

vircns Roxb. 497, 528 
virescens 528 

vitcllinum (Bl.) Dietr. 497, 
498, 514 

var. clovis Steen. ex 
Meijden 517 
wrayi King 497, 525 
Ximenia L. 1-6, 10 
americana L. 2, 5, 11 

var. americana 11, 12* 
borneensis Baill. 15 
loranlhifolia Span. 1 1 
olacioides W. & A. 47 
Xylosma leprusipes Clos 

Xyridaceae 4: 366 — 376, 

598; 5: 557; 9: 571 

Zanonia indica L. 253 
Zizyphus liitorea Teysm. ex 

Hassk. 1 1 
Zonalapollenites 338 
Zygophyllaceae 4: 64 

New York Botanical Garden Librar 

3 5185 00275 8033 




I •• 














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